Barbara Zimmermann

• Malick-Wahls, Sarah Lou; Devineau, Olivier; Zimmermann, Barbara & Tofastrud, Morten (2026). Finding common ground in commercial forests: Sheep grazing, timber production, and carnivore fences. Forest Ecology and Management. ISSN 0378-1127. 609. doi: 10.1016/j.foreco.2026.123663. Full text in Research Archive Show summary

  Sheep (Ovis aries Linnaeus, 1758) grazing has the potential for positive impacts on commercial forestry through weeding competing plants, or to be detrimental if browsing and trampling levels are high. In some regions of Norway, large, forested areas are fenced to minimize depredation risk from recovering carnivore populations on unsupervised sheep, but fences may alter the behavior of sheep leading to differential effects inside and outside of these fences. We estimated Norway spruce (Picea abies (Karst.) L.) sapling growth and frequency of damage in Inland and Trøndelag counties Norway, and related these to grazing intensity, fenced status, and competing vegetation along 70 transects within 8 fenced and 11 unfenced clearcuts during August 2024. We found that growth of terminal leaders decreased with increasing grazing intensity inside fences, whereas growth increased along with grazing intensity outside fences. Sheep grazing in our study had a positive effect on spruce sapling growth form by decreasing slenderness, measured by height-to-diameter ratio, both inside and outside fences. While less than three percent of saplings were browsed overall, browsing of saplings did increase at higher grazing intensities. Trampling was negligible on all transects. Our results suggest that sheep grazing is compatible with spruce regeneration and carnivore fencing as negative impacts from increasing amounts of sheep resulting in reduced growth of terminal leaders may have been offset by decreasing height-to-diameter ratios. Lower height-to-diameter ratios suggest more resilient saplings that could make them better able to withstand stressors. However, if sheep are allowed to reach high grazing intensities, this balance may not be achieved as browsing increases. With increasing demands on forested landscapes, integrating sheep grazing in carnivore-resistant fences on commercial forests presents an opportunity for multiple-use land management with minimal associated trade-offs.

• Beyene, Solomon Mulat; Naumov, Vladimir; Manton, Michael; Zimmermann, Barbara & Angelstam, Pär Krister (2026). Spatial variation in game species assemblages across a coastal-inland landscape gradient in Norway: associations with land cover and elevation and implications for landscape management. European Journal of Wildlife Research. ISSN 1612-4642. 72(3). doi: 10.1007/s10344-026-02095-7. Full text in Research Archive Show summary

  Game species are valuable resources in many regions and contribute to a range of ecosystem services, yet they are often studied and managed individually despite responding to similar environmental conditions. This study examined spatial variation in 10 game species across a coastal-inland gradient in Norway by (1) examining regional variation in relative abundance, and (2) analysing spatial patterns of game species assemblages in relation to land-cover composition and elevation. We used harvest density as a proxy for relative abundance, and analysed data using negative binomial regression, self-organising maps, and principal component analysis. Significant regional differences were observed for several species, including moose (Alces alces), red deer (Cervus elaphus), roe deer (Capreolus capreolus), black grouse (Lyrurus tetrix), mountain hare (Lepus timidus), and capercaillie (Tetrao urogallus). The relative harvest density of red fox (Vulpes vulpes), pine marten (Martes martes), rock ptarmigan (Lagopus muta), and willow grouse (Lagopus lagopus) did not differ significantly among regions. Spatial clustering identified five game species assemblages (red deer, willow grouse, moose-forest grouse, roe deer-moose, willow grouse–ptarmigan) defined by distinct combinations of land-cover types, elevation and harvest densities across municipalities. Assemblages were distributed across gradients from coastal semi-natural landscapes to inland forest–agricultural mosaics and from lowland to higher-elevation environments. Our findings show that game species assemblages vary systematically with landscape composition and elevation across the coastal–inland gradient. This highlights the value of an assemblage-based perspective for understanding spatial patterns and integrating game species management with land-use planning.

• Veenbrink, Wessel; Zimmermann, Barbara; Sand, Håkan; Wikenros, Camilla; Wabakken, Petter & Mayer, Martin (2026). Wolves on ice: habitat use and selection of water bodies by wolves during winter. Animal Behaviour. ISSN 0003-3472. 231. doi: 10.1016/j.anbehav.2025.123401. Full text in Research Archive Show summary

  Terrestrial mammals are in many ways connected to water bodies, although studies on habitat selection within anthropogenic landscapes often overlook the importance of water. The recolonization of grey wolves, Canis lupus, across parts of Europe, together with developments in GPS technology, has offered opportunities to study detailed spatial movements of this large carnivore in human-influenced landscapes. Here, we investigated wolf use, habitat selection and step length in relation to water bodies in the boreal forest during winter. We hypothesized that wolves select for larger water bodies during ice periods and use them as travel corridors for efficient movement, similar to forest roads. We used step selection analyses with data from 71 GPS-collared adult wolves in 44 wolf territories in Scandinavia to investigate habitat selection in relation to water bodies, time of day and season. The study included >50 000 GPS positions taken at 4 h intervals between 2001 and 2023. Wolves selected for lakes and rivers during ice periods, especially at night, with step length increasing when travelling on ice. Deep snow generally impeded wolf step length, but not more on frozen water than in other habitats. These results indicate that wolves utilize frozen water bodies as travel corridors for easier mobility during winter, possibly to reduce human-encounter risk. Our findings emphasize the adaptability of wolves and, more generally, contribute to our understanding of mammalian movement patterns and space use in the boreal zone.

• Auclair, Léa; Bernardi, Cecilia Di; Sand, Håkan; Åkesson, Mikael; Zimmermann, Barbara & Flagstad, Øystein [Show all 8 contributors for this article] (2025). Recruitment probability in a large carnivore: the role of biological and human‐related factors in early‐life. Oikos. ISSN 0030-1299. 2026(1). doi: 10.1002/oik.11422. Full text in Research Archive Show summary

  To reach reproduction, individuals must survive the juvenile stage, a critical period of low survival rates in large carnivores. Early‐life conditions during this stage can have lasting effects on survival, reproductive maturation, growth, physiology and behaviour. We assessed recruitment probability in Scandinavian wolves, i.e. the probability that a wolf reaches the reproductive stage and has pups surviving at least five months of age. To unravel human‐related and biological factors within the natal territory that could affect recruitment probability, we analysed life‐history data from 582 Scandinavian wolves Canis lupus identified by DNA as pups or juveniles in their birth territory. Factors considered included main prey density, road density, human density, and proximity to non‐breeding zones, as well as sex, inbreeding level and collaring. Among the 582 wolves analysed, 122 produced at least one surviving pup, corresponding to a recruitment probability of 0.21. Recruitment probability was more than twice as high (0.5) for juvenile wolves fitted with GPS‐collars compared to non‐collared individuals (0.22), and was positively correlated with human population density in the natal territory. We found no significant effects of other biologically or human‐related predictors. These results suggest that in this large carnivore population, managed below carrying capacity, individual recruitment probability is primarily influenced by human‐related factors, potentially reflecting poaching risk.

• Ausilio, Giorgia; Wikenros, Camilla; Sand, Håkan; Devineau, Olivier; Wabakken, Petter & Eriksen, Ane [Show all 10 contributors for this article] (2025). Contrasting risk patterns from human hunters and a large carnivore influence the habitat selection of shared prey. Oecologia. ISSN 0029-8549. 2017. doi: 10.1007/s00442-025-05742-z. Full text in Research Archive Show summary

  Spatial patterns of human hunting and predation risk are mediated by the physical landscape, with hunting risk often associated with different habitat features than those linked to large carnivores. Risk from hunters and carnivores also varies over time, and prey may adjust anti-predator strategies accordingly. We used GPS data from 17 female moose (Alces alces) tracked over two fall–winter seasons (2018–2020) in south-central Scandinavia to study moose habitat selection in response to diel and seasonal variation in risk from wolves (Canis lupus) and human hunters. Predation risk was quantified using spatial models based on known wolf and hunter kill-sites. We applied resource selection functions and Generalized Additive Mixed Models to model moose habitat selection in response to wolf and hunting risk. Moose avoided high-risk hunting areas during the day throughout the hunting season but relaxed this avoidance at night and after the season ended. In contrast, we found no evidence that moose adjusted habitat use in response to diel or seasonal variation in wolf predation risk. These results suggest that human hunting was the primary driver of moose habitat selection during the hunting season, becoming less relevant when hunting ceased. Our findings highlight the dominant role of human risk in shaping prey behaviour and the importance of accounting for hunting when evaluating predator–prey dynamics in human-dominated systems. An increased understanding of the risk effects arising from humans and large carnivores on prey can deepen our understanding of the ecological roles of predators and humans.

• Milleret, Cyril Pierre; Dupont, Pierre; Dey, Soumen; Brøseth, Henrik; Kindberg, Jonas & Turek, Daniel [Show all 11 contributors for this article] (2025). Map of death: Spatially explicit mortality of the grey wolf. Proceedings of the Royal Society of London. Biological Sciences. ISSN 0962-8452. 292(2053). doi: 10.1098/rspb.2025.0948. Full text in Research Archive Show summary

  Death is an inherently spatial process. It happens to someone, somewhere, but often remains undetected in nature. Death is also the primary means by which humans regulate wildlife populations. Using a novel analytical method that accounts for the cryptic nature of the fate of individuals and one of the world’s most comprehensive non-invasive genetic monitoring datasets, we were able to map cause-specific mortality of the entire Scandinavian grey wolf (Canis lupus) population despite the fact that most mortality events (mean = 65%; [95% CrI: 50.3–76.8%]) remained undetected. Our analysis revealed strong spatial variation in mortality with, for example, areas with a high risk of mortality linked with the current wolf management policies. Furthermore, we showed that the risk of legal mortality increased, while the risk of mortality due to causes other than legal mortality decreased with local wolf population density. This illustrates the complex interactions between spatial determinants and cause-specific mortality and therefore the importance of considering spatial variation when estimating mortality. Maps of mortality can inform wildlife management and conservation by capturing an elusive process in population dynamics as it unfolds in time and space.

• Sand, Håkan; Zimmermann, Barbara; Wabakken, Petter; Eriksen, Ane & Wikenros, Camilla (2025). Quantifying large carnivore predation relative to human harvest on moose in an intensively managed boreal ecosystem. Ecological Applications. ISSN 1051-0761. 35(1). doi: 10.1002/eap.70000. Full text in Research Archive Show summary

  The return of large carnivores to areas with strong anthropogenic impact often results in conflicts among different interest groups. One cause of conflict is that large carnivores compete with humans for wild game species. In Scandinavia, the recolonization of wolves (Canis lupus) and brown bears (Ursus arctos) has important ramifications for the harvest of an ungulate species with high economic and recreational value, the moose. We estimated wolf and brown bear predation rates on moose (Alces alces) relative to harvest, natural causes of death, and vehicle collisions within 20 wolf territories. We used data on multi-season kill rates of wolves and brown bears on moose combined with wolf territory sizes and estimates of the population density of brown bears and moose. Wolf predation rate on moose was not related to the density of moose, wolf pack size, nor kill rate but was positively related to wolf density and strongly negatively related to the abundance of moose within wolf territories. Estimated annual wolf and brown bear predation rates averaged 8.6% (range 2.8%–16.9%) and 2.3% (range 0%–12.7%) respectively, among wolf territories, whereas estimated annual harvest rates averaged 17.5% (range 8.1%–33.1%). In wolf territories with relatively high bear densities, the combined predation rates from wolves and brown bears exceeded harvest rates. Across wolf territories, harvest rates were not related to wolf predation rates or to the combined predation rates from wolves and brown bears, indicating that large carnivore predation and harvest were not compensatory to each other at this spatial level. The recolonization of these large carnivores in the Scandinavian boreal forest ecosystem may have small to significant consequences for the sustainable management of moose populations depending on the local conditions of both wolves, brown bears, and moose. Comparison of annual mortality rates for moose in our study in Scandinavia with corresponding data from areas with lower anthropogenic impact (Alaska) shows lower total mortality rates in Scandinavia. This likely results from a different age and sex composition of moose killed by wolves and brown bears versus harvest, in combination with a significant difference in the relative importance of these mortality factors between areas.

• Wikenros, Camilla; Zimmermann, Barbara; Månsson, Johan; Eriksen, Ane; Wabakken, Petter & Sand, Håkan (2025). Retrospective analyses to understand how wolf territory density impacts moose quotas, harvest and observation rate. European Journal of Wildlife Research. ISSN 1612-4642. 71. doi: 10.1007/s10344-025-01920-9. Full text in Research Archive Show summary

  The ever-growing human population along with the expansion of settlements and land use, and effective hunting methods increasingly influence wildlife populations. Knowledge of management responses to re-establishing large carnivores is important to understand the overall impact of humans on large carnivores and their prey populations. We examined the response of moose (Alces alces) harvest, but also population size and composition in relation to wolf (Canis lupus) territory density along a latitudinal gradient in two bordering countries in northern Europe. Moose harvest density was negatively related to wolf territory density with model estimates showing that harvest was 35% (Norway) to 39% (Sweden) lower in moose management units (MMUs) with average wolf territory density, compared to MMUs without wolves during the previous five years. The corresponding model estimates for moose observation rate was 21% lower in Sweden and 1% lower in Norway. In both countries, management actions were taken to reduce the total moose mortality (reduced harvest) as well as to maximize productivity in the population (reduced harvest of adult females) in response to increased wolf territory density. Annual changes in quotas were related to fulfilment of last year´s quota and wolf territory density in the previous two years. The annual change in harvest were affected by actual harvest the previous year and by set quotas, showing that harvest followed management plans. Abilities to adjust to new conditions is a key in wildlife management where conflicting societal objectives such as forestry, sustainable ungulate harvest yield, and carnivore conservation should be balanced.

• Mayer, Martin; Dyngeland, Cecilie; Wabakken, Petter; Tofastrud, Morten & Zimmermann, Barbara (2025). Patterns of cattle and sheep losses related to large carnivores and other causes in the outfields of Norway. Biological Conservation. ISSN 0006-3207. 305. doi: 10.1016/j.biocon.2025.111107. Full text in Research Archive Show summary

  Large carnivore populations have increased in many areas of Europe, resulting in socio-political conflicts, especially in areas with extensive outfield grazing systems. In Norway, the occurrence of large carnivores, especially brown bears (Ursus arctos) and grey wolves (Canis lupus), are largely separated from outfield grazing areas via carnivore management zones. However, the overall drivers of livestock losses in the outfields and the contribution of large carnivores are not well understood. Here, we investigated (1) correlates of cattle and sheep losses across organized grazing areas of Norway to evaluate which factors, including large carnivore densities, livestock density, the size of the grazing area, tick density, landscape characteristics, and climatic variables, are responsible for livestock losses in the outfields, and (2) spatio-temporal correlates of cattle losses suspected of being attacked by large carnivores. Based on our analyses investigating cattle and sheep losses in organized outfield grazing areas, we found no evidence that large carnivore densities correlated with cattle losses, but the proportion of sheep losses increased with increasing large carnivore densities, especially of bears and wolverines (Gulo gulo). Importantly, factors other than large carnivores, such as tick density and landscape composition, were related to higher probabilities of cattle losses. Our study provides important information concerning the overall causes of livestock losses in the outfields, indicating that large carnivores contribute little to total cattle losses.

• Forti, Alessandro; Lissillour, Pierre; Eriksen, Ane; Cerjak, Benoit; Campon, Carla & Motlova, Sona [Show all 8 contributors for this article] (2024). Carnivore exclosures to protect sheep affect the distribution of a wild cervid. Wildlife Biology. ISSN 0909-6396. doi: 10.1002/wlb3.01301. Full text in Research Archive Show summary

  With the recent recovery of large carnivores in Europe, preventive measures to protect livestock are on the rise. Fences that exclude carnivores from grazing areas have been proven as effective, but they can be costly as well as posing a barrier for wildlife. We studied the effect of exclosures of > 10 km2 to protect sheep Ovis aries on the distribution and density of moose Alces alces using fecal pellet group counts in two study areas in southeastern Norway. During the summer grazing season, the fences were powered. Outside of the grazing season, one exclosure remained fenced while the other fence was demounted. This quasi-experimental setting allowed us to investigate whether fences had a barrier effect for moose, and/or whether moose density was affected by interactions with sheep (competition or facilitation) or large carnivores (refuge hypothesis). During winter, moose pellet group density was about equal inside and outside of the exclosure with demounted wire strands, but less than half inside the permanently fenced exclosure compared to outside, indicating a potential fragmentation effect of the fence. During the grazing season, when wire strands were powered, moose pellet group density was equal or doubled inside as compared to outside both exclosures. Moose may have sought refuge from large carnivores inside the fences. Fecal pellet group densities of moose and sheep inside the fence were neither positively (facilitation) nor negatively (competition) correlated. However, moose used young forest, the most used habitat type by sheep, to a lesser extent inside than outside of the exclosures, maybe due to interference competition. Our study demonstrates that livestock protection fences can have an impact on more than the targeted wildlife species. To understand the mechanisms behind direct and indirect effects of fences, monitoring the movement and survival of individuals by means of GPS and camera traps would be needed.

• Spedener, Mélanie; Tofastrud, Morten; Austrheim, Gunnar & Zimmermann, Barbara (2024). A Grazer in a Browser's Habitat: Resource Selection of Foraging Cattle in Productive Boreal Forest. Rangeland Ecology & Management. ISSN 1550-7424. 93, p. 15–23. doi: 10.1016/j.rama.2023.12.004. Full text in Research Archive Show summary

  In Norway, cattle (Bos taurus) are released to large areas of boreal forest for summer grazing. To determine to what degree this practice challenges timber production and wildlife management, we need a better understanding of basic cattle ecology. What do cattle, typical grazers, feed on in a habitat typically used by browsers? We determined cattle's resource use and selection at three scales: habitat and microhabitat selection when foraging and diet selection. Boreal forest is dominated by grass-poor habitats, and despite their strong selection for grass-rich habitats, cows spent a lot of time in suboptimal habitats, like old bilberry (Vaccinium myrtillus) spruce (Picea abies) forest. However, they managed to find grass-rich habitat patches within those, selecting for patches with bentgrass (Agrostis spp.). Graminoids, mainly wavy hair-grass (Avenella spp.), tussock grass (Deschampsia), and true sedges (Carex spp.), made up the biggest part of the fecal samples. Woody plants, mainly willow (Salix spp.), Vaccinium spp., pine (Pinus spp.), and birch (Betula spp.), made up 9.4% ± 4.7% of the samples, a value lower than reported for other forest grazing cattle. Cattle avoided woody plants in their diet and selected for graminoids. They preferred deciduous over coniferous species. At these low stocking densities, the cows were grazers in this browser's habitat. Moreover, they selected rather on habitat and patch scale than for individual plants within a given habitat patch, considered typical for a grazer. Their grass-rich diet indicates little overlap with the diet of local wild ungulates, which are mostly browsers. Their aversion of spruce and selection for graminoids and deciduous trees indicates low conflict risk for browsing damages and even a beneficial weeding effect on trees planted for timber production.

• Nordli, Kristoffer; Walton, Zea Ann; Eriksen, Ane; Rogstad, Marius; Zimmermann, Barbara & Wikenros, Camilla [Show all 8 contributors for this article] (2024). Carcass provisioning and intra-guild risk avoidance between two sympatric large carnivores. Behavioral Ecology and Sociobiology. ISSN 0340-5443. 78. doi: 10.1007/s00265-023-03423-6. Full text in Research Archive Show summary

  Apex carnivores that rely primarily on predation play a central but complex role within scavenging ecology by potentially suppressing intra-guild competitors, but also facilitating them by providing a reliable supply of carrion. We investigated the competitive relationship between sympatric wolves (Canis lupus) and wolverines (Gulo gulo) in Norway across three seasons. We deployed remote cameras at fresh wolf kills (n = 29) and built Bayesian generalized linear mixed models (GLMMs) to explore the use of fresh wolf-killed prey by sympatric wolves and wolverines. Our results showed that wolves facilitated wolverines by providing scavenging opportunities. Biomass available from wolf kills was influenced by seasonal wolf prey preference and group size. Wolverines visited 100% of wolf kills in fall and winter, whereas only 18% in summer. We found that in winter, wolverines visited wolf kills 3.6 and 6.7 times more often than single wolves and wolf groups revisited their kills, and spent 10 and 25 times as much time at carcasses compared to single wolves and groups of wolves. Thus, wolverines played an important role in the depletion of wolf-killed prey, with potential effects on the scavenging behavior of other guild members. Understanding how globally threatened top predators may function as key species in scavenging processes is important to conservation as this may have community-wide cascading effects and support important ecosystem functions and services.

• Versluijs, Erik; Tofastrud, Morten; Hessle, Anna; Serrouya, Robert; Scasta, Derek & Wabakken, Petter [Show all 7 contributors for this article] (2024). Virtual fencing in remote boreal forests: performance of commercially available GPS collars for free-ranging cattle. Animal Biotelemetry. 12(1). doi: 10.1186/s40317-024-00389-8. Full text in Research Archive Show summary

  Background The use of virtual fencing in cattle farming is beneficial due to its flexibility, not fragmenting the landscape or restricting access like physical fences. Using GPS (Global Positioning System) technology, virtual fence units emit an audible signal and a low-energy electric shock when crossing a predefined border. In large remote grazing areas and complex terrains, where the performance of the GPS units can be affected by landscape structure, increased positioning errors can lead to unnecessary shocks to the animals leading to animal welfare concerns. This study aimed to explore factors affecting the GPS performance of commercially available virtual fence collars for cattle (NoFence©), both using static tests and mobile tests, i.e., when deployed on free-ranging cattle. Results The static tests revealed generally high fix success rates (% successful positioning attempts), and a lower success rate at four of 30 test locations was most likely due to a lack in GSM (Global System for Mobile communications) coverage. On average the GPS precision and accuracy errors were 3.3 m ± 2.5 SD and 4.6 m ± 3.2 SD, respectively. We found strong evidence that the GPS precision and accuracy errors increased errors under closed canopies. We also found evidence for an effect of the sky-view on the GPS performance, although at a lesser extent than canopy. The direction of the accuracy error in the Cartesian plane was not uniform, but biased, depending on the aspect of the test locations. With an average of 10.8 m ± 6.8 SD, the accuracy error of the mobile tests was more than double that of the static tests. Furthermore, we found evidence that more rugged landscapes resulted in higher GPS accuracy errors. However, the error from mobile tests was not affected by canopy cover, sky-view, or cattle behaviors. Conclusions This study showed that GPS performance can be negatively affected by landscape complexity, such as increased ruggedness and covered habitats, resulting in reduced virtual fence effectiveness and potential welfare concerns for cattle. These issues can be mitigated through proper pasture planning, such as avoiding rugged areas for the virtual fence border.

• Spedener, Mélanie; Mathisen, Karen Marie; Hauer, Josh; Hérault, Rémi; Austrheim, Gunnar & Tofastrud, Morten [Show all 7 contributors for this article] (2024). Effects of cattle grazing on young spruce trees in boreal production forest. Trees, Forests and People. 16. doi: 10.1016/j.tfp.2024.100596. Full text in Research Archive Show summary

  The compatibility of forest livestock grazing with timber production is disputed, as livestock can damage young trees through browsing and trampling. At the same time, livestock grazing might reduce the growth of competing vegetation and thereby enhance conifer forest regeneration. We investigated the effects of cattle (Bos taurus) on young spruce (Picea abies) trees in boreal production forest. We compared data from young spruce plantations within and outside of the grazing area, and during and off the cattle grazing season. Overall, stocking rates were 0.01 cows per hectare across the grazing areas. Cattle did not distribute equally, but concentrated their area use to young spruce plantations, leading to locally high densities during periods of the grazing season. Cattle removed vegetation competing with young spruce trees: Within the grazing area, the sward height of the field layer vegetation was lower, and young willow (Salix spec.) and downy birch (Betula pubescens) trees were browsed more than outside the grazing area. The annual height growth of spruce increased once the spruce grew above sward height. Spruce trees surrounded by many competing trees grew slower than those surrounded by few trees. Bent spruce trees grew slower than undamaged spruce trees. Yet, no difference in annual spruce growth was found between inside and outside the cattle grazing area. In contrast, we found a higher spruce mortality risk inside than outside the cattle grazing area. This mortality risk was very low and positively correlated with a given spruce tree's bark being damaged. Cattle did not increase the risk of damaged stem bark. This study did not succeed in pointing out the mechanism by which cattle might increase spruce mortality risk. Interestingly, the probability of being browsed and the probability of being bent were higher after winter than after summer, indicating that wild ungulates and snow might have a stronger effect on young spruce trees than summer grazing cattle at these stocking rates. In conclusion, cattle had minor effects on young spruce trees. This study suggests that at low stocking rates, forestry and livestock production in boreal production systems are compatible.

• Niccolai, Laura Jeanne; Devineau, Olivier; Thiel, Alexandra; Zimmermann, Barbara & Evans, Alina (2024). Connecting the dots: relationship between heart rate and overall dynamic body acceleration in free-ranging cattle. Conservation Physiology. 12(1). doi: 10.1093/conphys/coae085. Full text in Research Archive Show summary

  Monitoring physiological indicators including heart rate (HR) is crucial for managing animal welfare across diverse settings, from precision livestock farming to wildlife conservation. HR is a reliable indicator of energy expenditure and stress, yet the invasive nature of HR loggers limits their application in wild and free-ranging species. This study explores whether overall dynamic body acceleration (ODBA), measured with an external accelerometer, can serve as a less invasive proxy for HR. Using free-ranging cattle as a model species in Norway, we examined the relationship between ODBA and HR to assess how external accelerometry might indirectly reflect physiological states in settings that resemble wild conditions. Cattle provide an ideal model because they share some characteristics with wild herbivores, including exposure to diverse terrain and potential predation, whilst offering advantages for handling and sensor retrieval. Our findings showed that low ODBA values corresponded to static behaviours (e.g. standing, ruminating), where small movements caused HR spikes, whilst higher ODBA reflected dynamic activities (e.g. walking, foraging), with HR plateauing. This relationship suggests that ODBA can be used to approximate HR in environments where direct HR measurement is challenging. By using accelerometry to infer HR changes in free-ranging cattle, this study offers insights that could extend to wild species, offering a tool for conservationists to monitor and manage animal health and well-being less invasively.

• Benson, John F.; Keiter, David A.; Mahoney, Peter J.; Allen, Benjamin L.; Allen, Lee R. & Álvares, Francisco [Show all 102 contributors for this article] (2024). Intrinsic and environmental drivers of pairwise cohesion in wild Canis social groups. Ecology. ISSN 0012-9658. 106(1). doi: 10.1002/ecy.4492. Full text in Research Archive Show summary

  Animals within social groups respond to costs and benefits of sociality by adjusting the proportion of time they spend in close proximity to other individuals in the group (cohesion). Variation in cohesion between individuals, in turn, shapes important group-level processes such as subgroup formation and fission–fusion dynamics. Although critical to animal sociality, a comprehensive understanding of the factors influencing cohesion remains a gap in our knowledge of cooperative behavior in animals. We tracked 574 individuals from six species within the genus Canis in 15 countries on four continents with GPS telemetry to estimate the time that pairs of individuals within social groups spent in close proximity and test hypotheses regarding drivers of cohesion. Pairs of social canids (Canis spp.) varied widely in the proportion of time they spent together (5%–100%) during seasonal monitoring periods relative to both intrinsic characteristics and environmental conditions. The majority of our data came from three s

• Mayer, Martin; Furuhovde, Erlend; Nordli, Kristoffer; Ausilio, Giorgia; Wabakken, Petter & Eriksen, Ane [Show all 9 contributors for this article] (2024). Monitoring GPS-collared moose by ground versus drone approaches: efficiency and disturbance effects. Wildlife Biology. ISSN 0909-6396. doi: 10.1002/wlb3.01213. Full text in Research Archive Show summary

  Efficient wildlife management requires precise monitoring methods, for example to estimate population density, reproductive success, and survival. Here, we compared the efficiency of drone (equipped with a RGB camera) and ground approaches to detect and observe GPS-collared female moose Alces alces and their calves. We also quantified how drone (n = 42) and ground (n = 41) approaches affected moose behavior and space use (n = 24 individuals). The average time used for drone approaches was 17 min compared to 97 min for ground approaches, with drone detection probability being higher (95% of adult female moose and 88% of moose calves) compared to ground approaches (78% of adult females and 82% of calves). Drone detection success increased at lower drone altitudes (50–70 m). Adult female moose left the site in 35% of drone approaches (with > 40% of those moose becoming disturbed once the drone hovered < 50 m above ground) compared to 56% of ground approaches. We failed to find short-term effects (3 h after approaches) of drone approaches on moose space use, but moose moved > fourfold greater distances and used larger areas after ground approaches (compared to before the approaches had started). Similarly, longer-term (24 h before and after approaches) space use did not differ between drone approaches compared to days without known disturbance, but moose moved comparatively greater distances during days of ground approaches. In conclusion, we could show that drone approaches were highly efficient to detect adult moose and their calves in the boreal forest, being faster and less disturbing than ground approaches, making them a useful tool to monitor and study wildlife.

• Spedener, Mélanie; Valaker, Jenny Larsen; Helbert, Juliette; Schubert, Franziska Veronika; Mathisen, Karen Marie & Henriksen, Marie Vestergaard [Show all 9 contributors for this article] (2024). Effects of cattle and cervids on plants and flower-visiting insects in young spruce plantations. Forest Ecology and Management. ISSN 0378-1127. 572, p. 1–13. doi: 10.1016/j.foreco.2024.122258. Full text in Research Archive Show summary

  Livestock summer grazing in mountains and forests in Norway is generally considered beneficial to biodiversity. In this study we investigated if this is the case for cattle in boreal production forest. We collected biodiversity data on field layer vegetation, floral resources and flower-visiting insects in young spruce plantations that were planted 2–10 years ago. We picked young spruce plantations inside and outside well-established cattle grazing areas. On each plantation, we sampled fenced and unfenced plots (20 * 20 m each). This study design allowed us to investigate long-term effects of cattle grazing as well as short-term effects of excluding cervids only and short-term effects of excluding cervids and cattle. Long-term cattle grazing reduced the abundance of woody plants and reduced the abundance of flowers. Excluding cervids for two summers led to reduced height of woody plants (shrubs and heather) and to increased flower abundance. In contrast, excluding cervids and cattle for two summers led to increased height of graminoids, herbs and woody plants, to higher abundance of graminoids, higher flower abundance and higher abundance of flower-visiting insects. In conclusion, cattle affected the studied system in different ways and to a larger extent than native cervids. Our study shows that we must be careful when inferring results from cattle grazing studies on grasslands to forest ecosystems. As this study documents a negative effect of cattle on floral resources and flower-visiting insects, and we currently are facing a global pollination crisis, a careful consideration of the current practice of cattle grazing in boreal forest might be needed.

• Tofastrud, Morten; Møllevold, Ole Henrik Hammerstad & Zimmermann, Barbara (2024). Forage quality, road and stocking density influence the home range size of free-ranging beef cattle in the boreal forest. Applied Animal Behaviour Science. ISSN 0168-1591. 282. doi: 10.1016/j.applanim.2024.106455. Full text in Research Archive Show summary

  This study explores area use of beef cows (Bos taurus) free ranging in boreal production forest of south-eastern Norway. The forests in this region serve multiple purposes, with forestry being the predominant industry. Understanding cattle’s area use is crucial for implementing resource management strategies that balance ecological and economic sustainability of these forests. The aim of the study was to examine how extrinsic factors (stocking density, forage quality, road presence, and herd size) and intrinsic factors (reproductive status and breed) influence variations in seasonal and weekly home range sizes of beef suckler cows. The study also investigated changes in home range size over time. The study was conducted over three summers, using GPS collars programmed to record positions at 5 and 10-minute intervals on 75 cows of early and late maturing beef cattle breeds. The cows were allowed to roam freely in two areas, one with a high stocking density and the other with a low stocking density. The seasonal home ranges were found to be surprisingly large, averaging 3979 ± 344 ha and 2554 ± 246 ha ( ± SE) in the low and high stocking density areas, respectively. Both seasonal and weekly home range sizes positively correlated with the proportion of poor-quality vegetation types within the home range. In the low stocking density area, road density was negatively related to seasonal and weekly home range size. Weekly home ranges were largest in the low stocking density area and showed a slight increase throughout the grazing season. No correlation was found between home range size and cattle breed or reproductive status. In line with previous studies on area use by wild herbivores, vegetation quality significantly influenced the size of their home range. Therefore, vegetation mapping in grazing areas is an essential tool for livestock management in non-arable areas. The difference of home range size between the study areas may be due to factors other than stocking density. This finding should therefore be interpreted with caution, despite previous studies on wild herbivores confirming a negative correlation between population densities and home range size. Cows grazing in boreal forests use large areas and even larger when released to areas with poor forage quality, few roads and small stocking densities. This entails considerable movement throughout the grazing season, spanning across numerous habitats. This extensive area use can pose challenges for other stakeholders in the boreal forest and complicate the monitoring of cattle.

• Miltz, Cecilia; Eriksen, Ane; Wikenros, Camilla; Wabakken, Petter; Sand, Håkan & Zimmermann, Barbara (2024). Will future wind power development in Scandinavia have an impact on wolves? Wildlife Biology. ISSN 0909-6396. 2024(6). doi: 10.1002/wlb3.01250. Full text in Research Archive Show summary

  The global energy demand is growing, and the world is shifting towards using more renewable energy, like increased onshore wind power development. We used Global Positioning System (GPS) and Very High Frequency (VHF) location data from adult, territorial wolves Canis lupus in Scandinavia (Sweden and Norway; 1999–2021), to examine the potential for wind power development to affect wolf behavioural ecology. We examined the spatial overlap of areas proposed for wind power development with wolf territory activity centres prior to construction, to test to what extent overlap varies with season, time of day and social status (breeding versus non-breeding wolves). Measures of overlap were the distance between wolf activity centre points and nearest proposed wind turbine, the probability of proposed wind turbines being within the activity centre, and the density of proposed wind turbines within the activity centre. The wolf activity centre points were closer to sites of proposed turbines in early summer than in late winter and the density of proposed turbines in the activity centre was higher in early summer than in late winter. These findings probably result from an altitudinal shift in wolf area use between summer and winter. We also found that the probability for proposed turbines to be within the activity centre was higher for non-breeding than for breeding wolves during early summer, whereas it was higher for breeding compared to non-breeding wolves during late winter. This difference might be an effect of that breeding wolves have a restricted area use during the early summer season (denning period), resulting in a lower probability of turbines being inside their activity centre as compared to late winter. There was no clear pattern for other seasonal and social status differences. The results should be viewed as a starting point for further research and supplemented with before-after studies.

• Wikenros, Camilla; Sand, Håkan; Bernardi, Cecilia Di & Zimmermann, Barbara (2024). The role of predation, forestry and productivity in moose harvest at different spatial levels of management units. Wildlife Biology. ISSN 0909-6396. 2024(6). doi: 10.1002/wlb3.01248. Full text in Research Archive Show summary

  Management of ungulate populations to the desired density and/or demographic composition are challenged by contrasting aims of different stakeholders. For example, hunters may want to maximize hunting opportunities whereas commercial forest owners may want to minimize moose densities to mitigate browsing damage. In addition, the return of large predators such as wolves Canis lupus affects the possible harvest yield of ungulates and influences the population composition through their selection of specific age classes. The aim of this study was to gain a better understanding of factors related to the variation in moose Alces alces harvest. We used moose harvest statistics from the period 2012–2020, wolf annual monitoring data, annual brown bear Ursus arctos density, proportion of young forest per management unit, and proportion of agricultural land per management unit (index for productivity and roe deer Capreolus capreolus density) to explain variation in moose harvest across different management units at two spatial levels in two bordering countries, Sweden and Norway. The results showed variable responses in total harvest to changes in wolf territory density both at the regional and local management level. The proportion of young forest was correlated with both increased total harvest and proportion of calves. Increased proportion of agricultural land was linked to both increased total harvest and proportion of calves, likely due to that increased roe deer densities re-directed wolf predation from moose to roe deer, and an inverse relationship with brown bear density. Differences between countries may be due to differences in the management regime of moose, both in an historical and present perspective. Improved monitoring for individual hunting areas over time will be important for both the understanding of how different ungulate populations are affected by various factors and for the desired management of wildlife populations shared across borders.

• Ausilio, Giorgia; Sand, Håkan; Wikenros, Camilla; Aronsson, Malin; Milleret, Cyril Pierre & Nordli, Kristoffer [Show all 12 contributors for this article] (2023). Effects of large carnivores, hunter harvest, and weather on the mortality of moose calves in a partially migratory population. Wildlife Biology. ISSN 0909-6396. doi: 10.1002/wlb3.01179. Full text in Research Archive Show summary

  Survival of juvenile ungulates represents an important demographic parameter that influences population dynamics within ecosystems. In many ecological systems, the mortality of juvenile ungulates is influenced by various factors, including predation by large carnivores, human hunting activities and weather. While wolves Canis lupus are known to prey on moose Alces alces throughout all seasons, brown bears Ursus arctos primarily engage in predation during early summer, while human harvest primarily occurs in autumn and early winter. Hence, understanding the impacts of predation, harvest, and weather on the survival of juvenile moose is crucial for adaptive population management and the determination of sustainable harvest rates. To investigate the summer and autumn–winter survival of moose calves in relation to carnivore occurrence (wolf presence and bear density), summer habitat productivity, winter severity, human harvest, and migratory behaviour (migratory versus resident), we analysed data collected from 39 GPS-collared female moose in south-central Scandinavia. Our findings revealed significant interannual variation in summer survival rates, with areas with relatively higher bear densities exhibiting calf mortality rates twice as high as those in regions with low bear density. During the autumn–winter period, calf survival was lowest in the presence of wolves and deep snow, and it exhibited a negative correlation with the proportion of clearcuts and young forests within the mother's home range. Additionally, calf survival was negatively correlated with the risk of human hunting, and calves of stationary females displayed ten times higher survival rates compared to migratory individuals. Our study provides valuable insights into the survival of moose calves coexisting with two large carnivores and humans. Improving our understanding of the mechanisms causing calf survival to fluctuate has become increasingly important as many local moose populations in Scandinavia are declining and exposed to expanding predator populations, intense hunting pressure, and other threats associated with climate change.

• Nordli, Kristoffer; Wabakken, Petter; Eriksen, Ane; Sand, Håkan; Wikenros, Camilla & Maartmann, Erling [Show all 7 contributors for this article] (2023). Spatial and temporal cohesion of parents and offspring in a social large carnivore. Animal Behaviour. ISSN 0003-3472. 197, p. 155–167. doi: 10.1016/j.anbehav.2022.12.006. Full text in Research Archive Show summary

  Social organization in animals is a fundamental factor driving population dynamics and individual spatial distribution. Affiliation among kin is common in social groups, but kinship is no safeguard against intraspecific competition. Within social groups, the closest competitors are often related. In this study, we present 14 years of GPS-position movement data from 65 pairs (i.e. breeding pair, pup–adult breeder and pup–pup) of Scandinavian wolves, Canis lupus. We investigated social behaviour by examining spatial and temporal patterns of cohesion through winter to early summer (January to July) including two key reproductive periods, mating and birth, using mixed regressive nonlinear time series analysis (GAMM). During the mating period, breeding wolf pairs remained cohesive, while offspring increasingly dissociated. Offspring became even more solitary until the next birth period, when most permanently left their natal pack and territory. We suggest that the social organization of wolves may be modulated by intraspecific competition related to social aggression during the mating period and food competition around the birthing period. Early independence followed by immediate natal dispersal may be advantageous to both offspring and parents if the chances of finding food and a breeding partner and settling in a vacant area are high.

• Wikenros, Camilla; Bernardi, Cecilia Di; Zimmermann, Barbara; Åkesson, Mikael; Demski, Maike & Flagstad, Øystein [Show all 10 contributors for this article] (2023). Scavenging patterns of an inbred wolf population in a landscape with a pulse of human-provided carrion. Ecology and Evolution. 13(7). doi: 10.1002/ece3.10236. Full text in Research Archive Show summary

  Scavenging is an important part of food acquisition for many carnivore species that switch between scavenging and predation. In landscapes with anthropogenic impact, humans provide food that scavenging species can utilize. We quantified the magnitude of killing versus scavenging by gray wolves (Canis lupus) in Scandinavia where humans impact the ecosystem through hunter harvest, land use practices, and infrastructure. We investigated the cause of death of different animals utilized by wolves, and examined how the proportion of their consumption time spent scavenging was influenced by season, wolf social affiliation, level of inbreeding, density of moose (Alces alces) as their main prey, density of brown bear (Ursus arctos) as an intraguild competitor, and human density. We used data from 39 GPS-collared wolves covering 3198 study days (2001–2019),including 14,205 feeding locations within space–time clusters, and 1362 carcasses utilized by wolves. Most carcasses were wolf-killed (80.5%) while a small part had died from other natural causes (1.9%). The remaining had either anthropogenic mortality causes (4.7%), or the cause of death was unknown (12.9%). Time spent scavenging was higher during winter than during summer and autumn. Solitary wolves spent more time scavenging than pack-living individuals, likely because individual hunting success is lower than pack success. Scavenging time increased with the mean inbreeding coefficient of the adult wolves, possibly indicating that more inbred individuals resort to scavenging, which requires less body strength. There was weak evidence for competition between wolves and brown bears as well as a positive relationship between human density and time spent scavenging. This study shows how both intrinsic and extrinsic factors drive wolf scavenging behavior, and that despite a high level of inbreeding and access to carrion of anthropogenic origin, wolves mainly utilized their own kills. Canis lupus, consumption time, human density, inbreeding, intraguild competition, prey density, social affiliation

• Versluijs, Erik; Niccolai, Laura Jeanne; Spedener, Mélanie; Zimmermann, Barbara; Hessle, Anna & Tofastrud, Morten [Show all 8 contributors for this article] (2023). Classification of behaviors of free-ranging cattle using accelerometry signatures collected by virtual fence collars. Frontiers in Animal Science. 4, p. 1–13. doi: 10.3389/fanim.2023.1083272. Full text in Research Archive Show summary

  Precision farming technology, including GPS collars with biologging, has revolutionized remote livestock monitoring in extensive grazing systems. High resolution accelerometry can be used to infer the behavior of an animal. Previous behavioral classification studies using accelerometer data have focused on a few key behaviors and were mostly conducted in controlled situations. Here, we conducted behavioral observations of 38 beef cows (Hereford, Limousine, Charolais, Simmental/NRF/Hereford mix) free-ranging in rugged, forested areas, and fitted with a commercially available virtual fence collar (Nofence) containing a 10Hz tri-axial accelerometer. We used random forest models to calibrate data from the accelerometers on both commonly documented (e.g., feeding, resting, walking) and rarer (e.g., scratching, head butting, self-grooming) behaviors. Our goal was to assess pre-processing decisions including different running mean intervals (smoothing window of 1, 5, or 20 seconds), collar orientation and feature selection (orientation-dependent versus orientation-independent features). We identified the 10 most common behaviors exhibited by the cows. Models based only on orientation-independent features did not perform better than models based on orientation-dependent features, despite variation in how collars were attached (direction and tightness). Using a 20 seconds running mean and orientation-dependent features resulted in the highest model performance (model accuracy: 0.998, precision: 0.991, and recall: 0.989). We also used this model to add 11 rarer behaviors (each< 0.1% of the data; e.g. head butting, throwing head, self-grooming). These rarer behaviors were predicted with less accuracy because they were not observed at all for some individuals, but overall model performance remained high (accuracy, precision, recall >98%). Our study suggests that the accelerometers in the Nofence collars are suitable to identify the most common behaviors of free-ranging cattle. The results of this study could be used in future research for understanding cattle habitat selection in rugged forest ranges, herd dynamics, or responses to stressors such as carnivores, as well as to improve cattle management and welfare.

• Kirchner, Theresa Margret; Devineau, Olivier; Chimienti, Marianna; Thompson, Daniel P.; Crouse, John & Evans, Alina [Show all 8 contributors for this article] (2023). Predicting moose behaviors from tri-axial accelerometer data using a supervised classification algorithm. Animal Biotelemetry. 11(1). doi: 10.1186/s40317-023-00343-0. Full text in Research Archive Show summary

  Background Monitoring the behavior of wild animals in situ can improve our understanding of how their behavior is related to their habitat and affected by disturbances and changes in their environment. Moose (Alces alces) are keystone species in their boreal habitats, where they are facing environmental changes and disturbances from human activities. How these potential stressors can impact individuals and populations is unclear, in part due to our limited knowledge of the physiology and behavior of moose and how individuals can compensate for stress and disturbances they experience. We collected data from collar-mounted fine-scale tri-axial accelerometers deployed on captive moose in combination with detailed behavioral observations to train a random forest supervised classification algorithm to classify moose accelerometer data into discrete behaviors. To investigate the generalizability of our model to collared new individuals, we quantified the variation in classification performance among individuals. Results Our machine learning model successfully classified 3-s accelerometer data intervals from 12 Alaskan moose (A. a. gigas) and two European moose (A. a. alces) into seven behaviors comprising 97.6% of the 395 h of behavioral observations conducted in summer, fall and spring. Classification performance varied among behaviors and individuals and was generally dependent on sample size. Classification performance was highest for the most common behaviors lying with the head elevated, ruminating and foraging (precision and recall across all individuals between 0.74 and 0.90) comprising 79% of our data, and lower and more variable among individuals for the four less common behaviors lying with head down or tucked, standing, walking and running (precision and recall across all individuals between 0.28 and 0.79) comprising 21% of our data. Conclusions We demonstrate the use of animal-borne accelerometer data to distinguish among seven main behaviors of captive moose and discuss generalizability of the results to individuals in the wild. Our results can support future efforts to investigate the detailed behavior of collared wild moose, for example in the context of disturbance responses, time budgets and behavior-specific habitat selection.

• Neby, Magne; Andreassen, Harry Peter; Milleret, Cyril Pierre; Pedersen, Simen; Tamayo, Ana-Maria Peris & Sanchez, David Carricondo [Show all 8 contributors for this article] (2023). Small rodent monitoring at Birkebeiner Road, Norway. Biodiversity Data Journal. ISSN 1314-2836. 11. doi: 10.3897/BDJ.11.E105914. Full text in Research Archive Show summary

  Northern small mammal populations are renowned for their multi-annual population cycles. Population cycles are multi-faceted and have extensive impacts on the rest of the ecosystem. In 2011, we started a student-based research activity to monitor the variation of small rodent density along an elevation gradient following the Birkebeiner Road, in southeast Norway. Fieldwork was conducted by staff and students at the University campus Evenstad, Inland Norway University of Applied Sciences, which has a long history of researching cyclic population dynamics. The faculty has a strong focus on engaging students in all parts of the research activities, including data collection. Small rodents were monitored using a set of snap trap stations. Trapped animals were measured (e.g. body mass, body length, sex) and dissected to assess their reproductive status. We also characterised the vegetation at trapping sites.

• Versluijs, Erik; Eriksen, Ane; Fuchs, Boris; Wikenros, Camilla; Sand, Håkan & Wabakken, Petter [Show all 7 contributors for this article] (2022). Wolf Responses to Experimental Human Approaches Using High-Resolution Positioning Data. Frontiers in Ecology and Evolution. 10. doi: 10.3389/fevo.2022.792916. Full text in Research Archive Show summary

  Humans pose a major mortality risk to wolves. Hence, similar to how prey respond to predators, wolves can be expected to show anti-predator responses to humans. When exposed to a threat, animals may show a fight, flight, freeze or hide response. The type of response and the circumstances (e.g., distance and speed) at which the animal flees are useful parameters to describe the responses of wild animals to approaching humans. Increasing knowledge about behavioral responses of wolves toward humans might improve appropriate management and decrease conflicts related to fear of wolves. We did a pilot study by conducting 21 approach trials on seven GPS-collared wolves in four territories to investigate their responses to experimental human approaches. We found that wolves predominantly showed a flight response (N = 18), in a few cases the wolf did not flee (N = 3), but no wolves were seen or heard during trials. When wolves were downwind of the observer the flight initiation distance was significantly larger than when upwind, consistent with the hypothesis that conditions facilitating early detection would result in an earlier flight. Our hypothesis that early detection would result in less intense flights was not supported, as we found no correlation between flight initiation distances and speed, distance or straightness of the flight. Wolves in more concealed habitat had a shorter flight initiation distance or did not flee at all, suggesting that perceived risk might have been affected by horizontal visibility. Contrary to our expectation, resettling positions were less concealed (larger horizontal visibility) than the wolves’ initial site. Although our small number of study animals and trials does not allow for generalizations, this pilot study illustrates how standardized human approach trials with high-resolution GPS-data can be used to describe wolf responses at a local scale. In continuation, this method can be applied at larger spatial scales to compare wolf flight responses within and between populations and across anthropogenic gradients, thus increasing the knowledge of wolf behavior toward humans, and potentially improving coexistence with wolves across their range.

• Eriksen, Ane; Versluijs, Erik; Fuchs, Boris; Zimmermann, Barbara; Wabakken, Petter & Ordiz, Andres [Show all 15 contributors for this article] (2022). A Standardized Method for Experimental Human Approach Trials on Wild Wolves. Frontiers in Ecology and Evolution. 10. doi: 10.3389/fevo.2022.793307. Full text in Research Archive Show summary

  As wolves recolonize areas of Europe ranging from moderate to high anthropogenic impact, fear of wolves is a recurring source of conflict. Shared tools for evaluating wolf responses to humans, and comparing such responses across their range, can be valuable. Experiments in which humans approach wild wolves can increase our understanding of how wolves respond to humans, facilitating human-wolf coexistence. We have developed the first standardized protocol for evaluating wolf responses to approaching humans using high-resolution GPS data, and tested it on wild wolves. We present a field protocol for experimentally approaching GPS-collared wolves, a descriptive comparison of two statistical methods for detecting a measurable flight response, a tutorial for identifying wolf flight initiation and resettling positions, and an evaluation of the method when reducing GPS positioning frequency. The field protocol, a data collection form, and the tutorial with R code for extracting flight parameters are provided. This protocol will facilitate studies of wolf responses to approaching humans, applicable at a local, national, and international level. Data compiled in a standardized way from multiple study areas can be used to quantify the variation in wolf responses to humans within and between populations, and in relation to predictors such as social status, landscape factors, or human population density, and to establish a baseline distribution of wolf response patterns given a number of known predictors. The variation in wolf responses can be used to assess the degree to which results can be generalized to areas where GPS studies are not feasible, e.g., for predicting the range of likely wolf behaviors, assessing the likelihood of wolf-human encounters, and complementing existing tools for evaluating reports of bold wolves. Showing how wolves respond to human encounters should help demystify the behavior of wild wolves toward humans in their shared habitat.

• Smith, Adam F.; Ciuti, Simone; Shamovich, Dmitry; Fenchuk, Viktar; Zimmermann, Barbara & Heurich, Marco Dietmar (2022). Quiet islands in a world of fear: Wolves seek core zones of protected areas to escape human disturbance. Biological Conservation. ISSN 0006-3207. 276. doi: 10.1016/j.biocon.2022.109811. Full text in Research Archive Show summary

  The Anthropocene continuously escalates the challenges and threats faced by large carnivores in human-dominated landscapes. Given their unique conservation and management requirements, detailed insights into their behaviour in relation to human-induced risks are crucial to designing landscapes of coexistence for people and predators, containing key features such as nature conservation areas. Adaptations like selection or avoidance of areas at certain times reveal the dynamic perception of large carnivores to risk-landscapes and the trade-offs with potential resources. This study investigates these ideas by applying a habitat-selection approach to GPS collared wolves in Belovezhskaya Pushcha National Park (Belarus) to quantify fine-scale avoidance of human disturbance. Our results indicate that wolves selected strongly for areas of higher nature protection within the national park. They also avoided human disturbance from settlements and roads, but mediated their behaviour depending on the shifting risk throughout the day and night. They selected for more open areas during darkness. Seasonal and sex-based differences shifted avoidance, such as females being more avoidant of settlements. This suggests evidence of a successful anti-predator strategy against humans. Our study offers some of the most detailed and novel insights into wolf spatio-temporal avoidance of disturbance in the context of protected areas, which is imperative for fine-tuning management measures for continued coexistence and effective conservation of this functionally important species.

• Rodríguez-Recio, Mariano; Wikenros, Camilla; Zimmermann, Barbara & Sand, Håkan (2022). Rewilding by Wolf Recolonisation, Consequences for Ungulate Populations and Game Hunting. Biology (Basel). 11(2). doi: 10.3390/biology11020317. Full text in Research Archive Show summary

  Simple Summary: Humans extirpated the wolf Canis lupus from many regions of Europe. Today, the wolf is returning to many of these areas, and with it, people’s opposition due to its predatory habits on, among others, ungulate game species. Based on existing data on wolf prey selection, kill rates and territory size, we extrapolated the results from central Sweden and Poland to southern Sweden, where wolf recolonization has not yet occurred and conservation conflicts with hunters are expected. Thus, we calculated the proportion of moose Alces alces, roe deer Capreolus capreolus, red deer Cervus elaphus, fallow deer Dama dama and wild boar Sus scrofa that would be killed by wolves in the municipalities of southern Sweden if wolf recolonization occurs. We found that the current system of five ungulate species in southern Sweden could potentially support a wolf density two to four times higher than in the current wolf distribution in central Sweden, which are mainly inhabited by roe deer and moose. With this type of research, we can anticipate and work to ameliorate the social unrest and expected conservation conflicts that may arise once wolves or other large carnivore species recolonize areas of Europe that are returning to the wild. Abstract: The ongoing recolonisations of human-transformed environments in Europe by large carnivores like the wolf Canis lupus means that conservation conflicts could re-surface, among other reasons, due to predation on ungulate game species. We estimated the effect of wolves on ungulate species using data on wolf prey selection, kill rates and territory size to build a hypothetical case of future expansion. We extrapolated results on predation from the current wolf distribution in central Sweden and eastern Poland to the eventual wolf recolonisation of southern Sweden. We then calculated the proportion of five ungulate game species killed annually by wolves, and the ratio between the predicted annual predation by wolves given future colonization and the number of ungulates currently harvested by hunters. Results showed that wolf recolonization in southern Sweden would have a minor impact on the estimated population densities of red deer Cervus elaphus, fallow deer Dama dama and wild boar Sus scrofa, but is likely to lead to a significant reduction in human captures of moose Alces alces and roe deer Capreolus capreolus. The current five-ungulate species system in southern Sweden suggests a potential for two to four times higher wolf density than the two-ungulate species system in the northern part of their current distribution. Management and conservation of recolonizing large carnivores require a better understanding of the observed impact on game populations under similar ecological conditions to ameliorate conservation conflicts and achieve a paradigm of coexistence. Integrating these predictions into management is paramount to the current rewilding trend occurring in many areas of Europe or North America.

• Ausilio, Giorgia; Wikenros, Camilla; Sand, Håkan; Wabakken, Petter; Eriksen, Ane & Zimmermann, Barbara (2022). Environmental and anthropogenic features mediate risk from human hunters and wolves for moose. Ecosphere. 13(12). doi: 10.1002/ecs2.4323. Full text in Research Archive Show summary

  Landscape characteristics, seasonal changes in the environment, and daylight conditions influence space use and detection of prey and predators, resulting in spatiotemporal patterns of predation risk for the prey. When predators have different hunting modes, the combined effects of multiple predators are mediated by the physical landscape and can result in overlapping or contrasting patterns of predation risk. Humans have become super-predators in many anthropogenic landscapes by harvesting game species and competing with large carnivores for prey. Here, we used the locations of wolf (Canis lupus)-killed and hunter-killed moose (Alces alces) in south-central Scandinavia to investigate whether environmental and anthropogenic features influenced where wolves and hunters killed moose. We predicted that the combined effects of wolves and hunters would result in contrasting spatial risk patterns due to differences in hunting modes. We expected these contrasting spatial risk patterns also to differ temporally. During the hunting season, the probability of a wolf kill increased with distance to bogs, whereas it decreased with increasing building density and distance to clearcuts and young forests. After the hunting season, the probability of a wolf kill increased with increasing terrain ruggedness and decreased with increasing building density, distance to main roads, and distance to clearcuts and young forests. The probability of a hunter kill was highest closer to bogs, main and secondary roads, in less rugged terrain and in areas with lower building density. Hunters killed all moose during the day, whereas wolves killed most moose at night during and after the hunting season. Our findings suggest that environmental and anthropogenic features mediate hunting and wolf predation risk. Additionally, we found that hunter- and wolf-killed moose exhibited contrasting spatial associations to landscape features, most likely due to the different hunting modes displayed by hunters and wolves. However, wolf predation and hunting risks also contrasted over time since wolves killed mostly at night and hunters were restricted to hunting during daytime and during the hunting season. This temporal segregation in risk might therefore suggest that moose could minimize risk exposure by taking advantage of spatiotemporally vacant hunting domains.

• Loosen, Anne Elizabeth; Devineau, Olivier; Zimmermann, Barbara & Mathisen, Karen Marie (2022). The importance of evaluating standard monitoring methods: Observer bias and detection probabilities for moose pellet group surveys. PLOS ONE. 17(7). doi: 10.1371/journal.pone.0268710. Full text in Research Archive Show summary

  Counting is not always a simple exercise. Specimens can be misidentified or not detected when they are present, giving rise to unidentified sources of error. Deer pellet group counts are a common method to monitor abundance, density, and population trend. Yet, detection errors and observer bias could introduce error into sometimes very large (spatially, temporally) datasets. For example, in Scandinavia, moose (Alces alces) pellet group counts are conducted by volunteer hunters and students, but it is unknown how much uncertainty observer error introduces into these datasets. Our objectives were to 1) estimate the detection probability of moose pellet groups; 2) identify the primary variables leading to detection errors including prior observer experience; and 3) compare density estimates using single and double observer counts. We selected a subset of single observer plots from a long-term monitoring project to be conducted as dependent double observer surveys, where primary and secondary observers worked simultaneously in the field. We did this to quantify detection errors for moose pellet groups, which were previously unknown in Scandinavia, and to identify covariates which introduced variation into our estimates. Our study area was in the boreal forests of southern Norway where we had a nested grid of 100-m2 plots that we surveyed each spring. Our observers were primarily inexperienced. We found that when pellet groups were detected by the primary observer, the secondary observer saw additional pellet groups 42% of the time. We found search time was the primary covariate influencing detection. We also found density estimates from double observer counts were 1.4 times higher than single observer counts, for the same plots. This density underestimation from single observer surveys could have consequences to managers, who sometimes use pellet counts to set harvest quotas. We recommend specific steps to improve future moose pellet counts.

• Tallian, Aimee Grace; Ordiz, Andres; Metz, Matthew C.; Zimmermann, Barbara; Wikenros, Camilla & Smith, Douglas W. [Show all 11 contributors for this article] (2021). Of wolves and bears: Seasonal drivers of interference and exploitation competition between apex predators. Ecological Monographs. ISSN 0012-9615. 92(2). doi: 10.1002/ecm.1498. Full text in Research Archive Show summary

  Competition between apex predators can alter the strength of top-down forcing, yet we know little about the behavioral mechanisms that drive competition in multipredator ecosystems. Interactions between predators can be synergistic (facilitative) or antagonistic (inhibitive), both of which are widespread in nature, vary in strength between species and across space and time, and affect predation patterns and predator–prey dynamics. Recent research has suggested that gray wolf (Canis lupus) kill rates decrease where they are sympatric with brown bears (Ursus arctos), however, the mechanisms behind this pattern remain unknown. We used data from two long-term research projects in Scandinavia (Europe) and Yellowstone National Park (North America) to test the role of interference and exploitation competition from bears on wolf predatory behavior, where altered wolf handling and search time of prey in the presence of bears are indicative of interference and exploitation competition, respectively. Our results suggest the mechanisms driving competition between bears and wolves were dependent on the season and study system. During spring in Scandinavia, interference competition was the primary mechanism driving decreased kill rates for wolves sympatric with bears; handling time increased, but search time did not. In summer, however, when both bear and wolf predation focused on neonate moose, the behavioral mechanism switched to exploitation competition; search time increased, but handling time did not. Alternartively, interference competition did affect wolf predation dynamics in Yellowstone during summer, where wolves prey more evenly on neonate and adult ungulates. Here, bear presence at a carcass increased the amount of time wolves spent at carcasses of all sizes and wolf handling time for small prey, but decreased handling time for the largest prey. Wolves facilitate scavenging opportunities for bears, however, bears alter wolf predatory behavior via multiple pathways and are primarily antagonistic to wolves. Our study helps to clarify the behavioral mechanisms driving competition between apex predators, illustrating how interspecific interactions can manifest into population-level predation patterns.

• Wikenros, Camilla; Gicquel, Morgane; Zimmermann, Barbara; Flagstad, Øystein & Åkesson, Mikael (2021). Age at first reproduction in wolves: Different patterns of density dependence for females and males. Proceedings of the Royal Society of London. Biological Sciences. ISSN 0962-8452. 288(1948). doi: 10.1098/rspb.2021.0207. Full text in Research Archive Show summary

  Age at first reproduction constitutes a key life-history trait in animals and is evolutionarily shaped by fitness benefits and costs of delayed versus early reproduction. The understanding of how intrinsic and extrinsic changes affects age at first reproduction is crucial for conservation and management of threatened species because of its demographic effects on population growth and generation time. For a period of 40 years in the Scandinavian wolf (Canis lupus) population, including the recolonization phase, we estimated age at first successful reproduction (pup survival to at least three weeks of age) and examined how the variation among individuals was explained by sex, population size (from 1 to 74 packs), primiparous or multiparous origin, reproductive experience of the partner and inbreeding. Median age at first reproduction was 3 years for females (n = 60) and 2 years for males (n = 74), and ranged between 1 and 8–10 years of age (n = 297). Female age at first reproduction decreased with increasing population size, and increased with higher levels of inbreeding. The probability for males to reproduce later first decreased, reaching its minimum when the number of territories approached 40–60, and then increased with increasing population size. Inbreeding for males and reproductive experience of parents and partners for both sexes had overall weak effects on age at first reproduction. These results allow for more accurate parameter estimates when modelling population dynamics for management and conservation of small and vulnerable wolf populations, and show how humans through legal harvest and illegal hunting influence an important life-history trait like age at first reproduction. ecology, age at first birth, age at maturity, Allee effect, Canis lupus, primiparity, inbreeding

• Loosen, Anne Elizabeth; Devineau, Olivier; Skarpe, Christina; Zimmermann, Barbara; Cromsigt, Joris & Mathisen, Karen Marie (2021). Ungulate-adapted forestry shows promise for alleviating pine browsing damage. Forest Ecology and Management. ISSN 0378-1127. 482(118808). doi: 10.1016/j.foreco.2020.118808. Full text in Research Archive Show summary

  High densities of ungulates can increase human-wildlife conflicts. Where forestry is an important economy, intensive browsing can lead to browsing damage, resulting in volume losses, poor stand regeneration, and reduced timber quality. The forestry industry thus looks for practical, long-term measures to mitigate browsing damage. We tested the effect of two mitigation measures on moose (Alces alces) browsing behaviour and damage to Scots pine (Pinus sylvetris): (1) ungulate-adapted slash piles (i.e., palatable species only) created during felling to increase short-term food availability and (2) intensified soil scarification to increase long-term food availability (collectively, ‘ungulate-adapted forestry’). Our study occurred in southern Norway where we established fixed vegetation and moose faecal pellet plots at varying distances from conventional and ungulate-adapted slash piles and scarified stands. We evaluated the effects of ungulate-adapted slash piles and intensified scarification on the density of undamaged Scots pine, moose bite diameters, browsing pressure, and moose faecal pellet density. To assess the effect of spatial scale, we created 250 m, 500 m, and 1000-m radius buffers centered on each plot. We found that ungulate-adapted logging near our plots increased the density of undamaged pines, as compared to no and conventional logging. We found that logging in general led to smaller bite diameters. We also found that plots near conventional logging had higher browsing pressure, whereas browsing pressure near ungulate-adapted logging was similar to unlogged stands. For scarification, density of undamaged pine increased when the ungulate-adapted stand aged whereas undamaged pine decreased as conventional scarification stands aged. Browsing pressure showed a response at the smallest spatial scale only for ungulate-adapted scarification. Peak moose habitat use near conventional and ungulate-adapted scarified stands differed by stand age and distance from scarification. The overall effects of ungulate-adapted forestry were most pronounced at the smallest spatial scale (250 m). Our results support ‘ungulate-adapted’ forestry as a practical solution to mitigate browsing damage but uncertainty in some of our estimates suggest further research on the area treated is needed.

• Tallian, Aimee Grace; Ordiz, Andres; Zimmermann, Barbara; Sand, Håkan; Wikenros, Camilla & Wabakken, Petter [Show all 8 contributors for this article] (2021). The return of large carnivores: Using hunter observation data to understand the role of predators on ungulate populations. Global Ecology and Conservation. 27. doi: 10.1016/j.gecco.2021.e01587. Full text in Research Archive Show summary

  Large carnivores play a key ecological role in nature, yet quantifying the effects of predation at large spatiotemporal scales remains challenging. Wolves and brown bears have recovered in Sweden, where they share the same staple prey, moose. This ecosystem is representative of the Eurasian boreal realm, and makes an interesting case study for exploring the use of hunter-collected observation data to understand the ecological effects of large carnivore recovery in human-dominated ecosystems. We used over 50 million hours of moose ob-servation data collected by “citizen scientists” (i.e., moose hunters) between 2000 and 2017 to evaluate the role of recovering bear and wolf populations on an important ungulate vital rate, calf/cow ratio (the number of observed calves per female in fall). Calf/cow ratios were negatively correlated with both wolf and bear densities, suggesting that summer calf survival decreased via predation. Calf/cow ratios decreased by 7% and 17% in the northern and southern areas where bears and wolves were allopatric, respectively. Where wolves and bears were sympatric, the effect of predator densities was additive; calf/cow ratios decreased by 18%. However, both calf/cow ratios and moose densities declined over the last ~20 years across Sweden, including areas where wolves and bears were absent. While recolonizing large carnivores affected ungulate vital rates in Sweden’s human-dominated landscape, they were likely not the primary driver of long-term moose population trends. Our results re-inforce that citizen-collected data are a useful wildlife monitoring tool that help understand ecological processes, including the effects of recovering carnivores on prey populations. Brown bear Calf/cow ratio Carnivore recovery Predator-prey interactions Wolf

• Loosen, Anne Elizabeth; Devineau, Olivier; Zimmermann, Barbara; Cromsigt, Joris P. G. M.; Pfeffer, Sabine & Skarpe, Christina [Show all 7 contributors for this article] (2021). Roads, forestry, and wolves interact to drive moose browsing behavior in Scandinavia. Ecosphere. 12(1). doi: 10.1002/ecs2.3358. Full text in Research Archive Show summary

  As wild ungulate densities increase across Europe and North America, plant–herbivore interactions are increasingly important from ecological and economic perspectives. These interactions are particularly significant where agriculture and forestry occur and where intensive grazing and browsing by wild ungulates can result in economic losses to growing crops and trees. We studied plant–herbivore interactions in a moose (Alces alces)-dominant system where forestry is a primary economy, the primary and secondary road networks are extensive, and wolves (Canis lupus) are recolonizing. Wolves and humans use low-traffic, secondary roads, yet roadsides provide high-quality and quantity browse for moose. Foraging theory predicts that moose will respond to riskier landscapes by selecting habitats that reduce predation risk, sacrificing feeding time or food quality. As food becomes limiting, however, animals will accept higher predation risk in search of food. We predicted that road avoidance behavior would be strongest within wolf territories. In areas without wolves, moose should select roadsides for their high forage availability. To test these predictions, we measured moose browsing and counted pellet groups as a proxy for habitat use each spring in Norway and Sweden between 2008 and 2018, in areas with and without wolves and at different distances from primary and secondary roads. We used generalized linear mixed models to evaluate drivers of the probability of browsing occurrence and browsing pressure. We found that browsing occurrence increased closer to secondary roads but decreased closer to primary roads. We also found browsing patterns to vary among tree species. For Scots pine (Pinus sylvestris), the browsing occurrence was two times higher in young forests relative to non-young forests and decreased further from secondary roads. Wolf territory presence and probability had neutral or positive effect on browsing occurrence and pressure for all species. However, wolf territory presence had negative effects on browsing occurrence and pressure when interacting with secondary roads, young forest, or snow cover. We showed that roads can influence browsing patterns in Norway and Sweden. However, further research is needed, particularly in the face of continued infrastructure development in Scandinavia.

• Sebastián-González, Esther; Morales-Reyes, Zebensui; Botella, Francisco; Naves-Alegre, Lara; Pérez-García, Juan M. & Mateo-Tomás, Patricia [Show all 59 contributors for this article] (2021). Functional traits driving species role in the structure of terrestrial vertebrate scavenger networks. Ecology. ISSN 0012-9658. 102(12). doi: 10.1002/ecy.3519. Full text in Research Archive Show summary

  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

• Veen, Bert van der; Mattisson, Jenny; Zimmermann, Barbara; Odden, John & Persson, Jens (2020). Refrigeration or anti-theft? Food-caching behavior of Wolverines (Gulo gulo) in Scandinavia. Behavioral Ecology and Sociobiology. ISSN 0340-5443. 74(5). doi: 10.1007/s00265-020-2823-4. Full text in Research Archive Show summary

  Food-caching animals can gain nutritional advantages by buffering seasonality in food availability, especially during times of scarcity. The wolverine (Gulo gulo) is a facultative predator that occupies environments of low productivity. As an adaptation to fluctuating Food availability, wolverines cache perishable food in snow, boulders, and bogs for short- and long-term storage. We studied caching behavior of 38 GPS-collared wolverines in four study areas in Scandinavia. By investigating clusters of GPS locations, we identified a total of 303 food caches from 17 male and 21 female wolverines.Wolverines cached food all year around, from both scavenging and predation events, and spaced their caches widely within their home range.Wolverines cached food items on average 1.1 km from the food source andmade between 1 and 6 caches per source.Wolverines cached closer to the source when scavenging carcasses killed by other large carnivores; this might be a strategy to optimize food gain when under pressure of interspecific competition.When caching, wolverines selected for steep and rugged terrain in unproductive habitat types or in forest, indicating a preference for less-exposed sites that can provide cold storage and/or protection against pilferage. The observed year-round investment in caching by Wolverines underlines the importance of food predictability for survival and reproductive success in this species. Increasing temperatures as a consequence of climate change may provide newchallenges for wolverines by negatively affecting the preservation of cached food and by increasing competition from pilferers that benefit from awarmer climate. It is however still not fully understood which consequences this may have for the demography and behavior of the wolverine.

• Ordiz, Andres; Milleret, Cyril; Uzal, Antonio; Zimmermann, Barbara; Wabakken, Petter & Wikenros, Camilla [Show all 9 contributors for this article] (2020). Individual Variation in Predatory Behavior, Scavenging and Seasonal Prey Availability as Potential Drivers of Coexistence between Wolves and Bears. Diversity. 12(9). doi: 10.3390/d12090356. Full text in Research Archive Show summary

  Several large carnivore populations are recovering former ranges, and it is important to understand interspecific interactions between overlapping species. In Scandinavia, recent research has reported that brown bear presence influences gray wolf habitat selection and kill rates. Here, we characterized the temporal use of a common prey resource by sympatric wolves and bears and described individual and seasonal variation in their direct and/or indirect interactions. Most bear–wolf interactions were indirect, via bear scavenging of wolf kills. Bears used >50% of wolf kills, whereas we did not record any wolf visit at bear kills. Adult and subadult bears visited wolf kills, but female bears with cubs of the year, the most vulnerable age class to conspecifics and other predators, did not. Wolf and bear kill rates peaked in early summer, when both targeted neonate moose calves, which coincided with a reduction in bear scavenging rate. Some bears were highly predatory and some did not kill any calf. Individual and age-class variation (in bear predation and scavenging patterns) and seasonality (in bear scavenging patterns and main prey availability of both wolves and bears) could mediate coexistence of these apex predators. Similar processes likely occur in other ecosystems with varying carnivore assemblages. apex predators; bear; interspecific interactions; moose; predation; scavenging; wolf

• Wikenros, Camilla; Sand, Håkan; Månsson, Johan; Maartmann, Erling; Eriksen, Ane & Wabakken, Petter [Show all 7 contributors for this article] (2020). Impact of a recolonizing, cross‑border carnivore population on ungulate harvest in Scandinavia. Scientific Reports. 10(1), p. 1–11. doi: 10.1038/s41598-020-78585-8. Full text in Research Archive Show summary

  Predation from large carnivores and human harvest are the two main mortality factors afecting the dynamics of many ungulate populations. We examined long-term moose (Alces alces) harvest data from two countries that share cross-border populations of wolves (Canis lupus) and their main prey moose. We tested how a spatial gradient of increasing wolf territory density afected moose harvest density and age and sex composition of the harvested animals (n= 549,310), along a latitudinal gradient during 1995–2017. In areas containing average-sized wolf territories, harvest density was on average 37% (Norway) and 51% (Sweden) lower than in areas without wolves. In Sweden, calves made up a higher proportion of the moose harvest than in Norway, and this proportion was reduced with increased wolf territory density, while it increased in Norway. The proportion of females in the adult harvest was more strongly reduced in Sweden than in Norway as a response to increased wolf territory density. Moose management in both countries performed actions aimed to increase productivity in the moose population, in order to compensate for the increased mortality caused by wolves. These management actions are empirical examples of an adaptive management in response to the return of large carnivores.

• Sebastián-González, Esther; Morales-Reyes, Zebensui; Botella, Francisco; Naves-Alegre, Lara; Pérez-García, Juan M. & Mateo-Tomás, Patricia [Show all 57 contributors for this article] (2020). Network structure of vertebrate scavenger assemblages at the global scale: drivers and ecosystem functioning implications. Ecography. ISSN 0906-7590. 43(8), p. 1143–1155. doi: 10.1111/ecog.05083. Full text in Research Archive Show summary

  The organization of ecological assemblages has important implications for ecosystem functioning, but little is known about how scavenger communities organize at the global scale. Here, we test four hypotheses on the factors affecting the network structure of terrestrial vertebrate scavenger assemblages and its implications on ecosystem functioning. We expect scavenger assemblages to be more nested (i.e. structured): 1) in species-rich and productive regions, as nestedness has been linked to high competition for carrion resources, and 2) regions with low human impact, because the most efficient carrion consumers that promote nestedness are large vertebrate scavengers, which are especially sensitive to human persecution. 3) We also expect climatic conditions to affect assemblage structure, because some scavenger assemblages have been shown to be more nested in colder months. Finally, 4) we expect more organized assemblages to be more efficient in the consumption of the resource. We first analyzed the relationship between the nestedness of the scavenger assemblages and climatic variables (i.e. temperature, precipitation, temperature variability and precipitation variability), ecosystem productivity and biomass (i.e. NDVI) and degree of human impact (i.e. human footprint) using 53 study sites in 22 countries across five continents. Then, we related structure (i.e. nestedness) with its function (i.e. carrion consumption rate). We found a more nested structure for scavenger assemblages in regions with higher NDVI values and lower human footprint. Moreover, more organized assemblages were more efficient in the consumption of carrion. However, our results did not support the prediction that the structure of the scavenger assemblages is directly related to climate. Our findings suggest that the nested structure of vertebrate scavenger assemblages affects its functionality and is driven by anthropogenic disturbance and ecosystem productivity worldwide. Disarray of scavenger assemblage structure by anthropogenic disturbance may lead to decreases in functionality of the terrestrial ecosystems via loss of key species and trophic facilitation processes.

• Tofastrud, Morten; Hessle, Anna; Rekdal, Yngve & Zimmermann, Barbara (2020). Weight gain of free-ranging beef cattle grazing in the boreal forest of south-eastern Norway. Livestock Science. ISSN 1871-1413. 233. doi: 10.1016/j.livsci.2020.103955. Full text in Research Archive Show summary

  Young forest stands and clearcuts in the boreal forest created by modern forestry practices along with meadows of abandoned summer farms may contribute as feeding areas for beef cattle. The patchy distribution and varying quality and diversity of forage on such unimproved lands may affect cattle productivity. Weight gain of 336 beef cows and 270 calves free-ranging during three summer grazing seasons was monitored in boreal forests of southeastern Norway, stocked at either high (0.16 cows ha-1) and low (0.04 cows ha-1) stocking densities. We used linear mixed effect models for assessing intrinsic correlates of weight gain in cows and calves in the two areas. Habitat use and home range size of a subsample of 53 cows were monitored by using GPS collars programmed to log locations at 5 min. intervals during the grazing season. Additional extrinsic correlates of weight gain for the subsampled cows using a linear mixed model were also tested. Average weight gain of beef cows grazing at the low stocking density was positive among cows of early maturing breeds (represented by Hereford) gaining 24 ± 2.8 kg ( ± SE), while cows of late maturing breeds (mainly represented by Charolais) had an average weight loss of 9 ± 8.4 kg. The average weight gain was negative for beef cows of both early (Herefords) and late maturing breeds (mainly represented by Charolais but also Limousin and Simmental) at the high stocking density. Within both breed groups, there was a negative relationship between breed-specific average weight of cows at turnout and weight gain during the grazing period, while a prolonged grazing period was slightly positively related to weight gain. There was no relationship between weight gain and home range size and proportion of grazing habitat for the 53 cows fitted with GPS collars. Higher weight gains in calves of the low compared to the high stocking density area was found. However, there was no breed effect of weight gain in calves. Across study areas, spring-born suckler calves gained more weight than autumn-born calves (92 ± 1.7 kg vs. 65 ± 4.4 kg). Also, there were higher weight gains for springborn bull-calves than spring-born heifers (100 ± 2.4 kg vs. 94 ± 2.2 kg). Overall, the results indicate that it is possible to achieve acceptable weight gains for cattle grazing coniferous forest by finding breeds suitable for these extensive areas and stocking at moderate densities.

• Ordiz, Andres; Uzal, Antonio; Milleret, Cyril Pierre; Sanz-Perez, Ana; Zimmermann, Barbara & Wikenros, Camilla [Show all 10 contributors for this article] (2020). Wolf habitat selection when sympatric or allopatric with brown bears in Scandinavia. Scientific Reports. 10. doi: 10.1038/s41598-020-66626-1. Full text in Research Archive Show summary

  Habitat selection of animals depends on factors such as food availability, landscape features, and intraand 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.

• Sanchez, David Carricondo; Zimmermann, Barbara; Wabakken, Petter; Eriksen, Ane; Milleret, Cyril Pierre & Ordiz, Andres [Show all 8 contributors for this article] (2020). Wolves at the door? Factors influencing the individual behavior of wolves in relation to anthropogenic features. Biological Conservation. ISSN 0006-3207. 244. doi: 10.1016/j.biocon.2020.108514. Full text in Research Archive Show summary

  The recovery of large carnivores in human-dominated landscapes comes with challenges. In general, large carnivores avoid humans and their activities, and human avoidance favors coexistence, but individual variation in large carnivore behavior may occur. The detection of individuals close to human settlements or roads can trigger fear in local communities and in turn demand management actions. Understanding the sources of individual variation in carnivore behavior towards human features is relevant and timely for ecology and conservation. We studied the movement behavior of 52 adult established wolves ((Canis lupus), 44 wolf pairs) with GPS-collars over two decades in Scandinavia in relation to settlements, buildings, and roads. We fit fine-scale movement data to individual step selection functions to depict the movement decisions of wolves while travelling, and then used weighted linear mixed models to identify factors associated with potential individual pair deviations from the general behavioral patterns. Wolves consistently avoided human settlements and main roads, with little individual variation. Indeed, after correcting for season, time of the day, and latitude, there was little variability in habitat selection among wolf pairs, demonstrating that all wolf pairs had similar movement pattern and generally avoided human features of the landscape. Wolf avoidance of human features was lower at higher latitudes particularly in winter, likely due to seasonal prey migration. Although occasional sightings of carnivores or their tracks near human features do occur, they do not necessarily require management intervention. Communication of scientific findings on carnivore behavior to the public should suffice in most cases.

• Milleret, Cyril; Ordiz, Andres; Perez, Ana Sanz; Uzal, Antonio; Sanchez, David Carricondo & Eriksen, Ane [Show all 11 contributors for this article] (2019). Testing the influence of habitat experienced during the natal phase on habitat selection later in life in Scandinavian wolves. Scientific Reports. 9. doi: 10.1038/s41598-019-42835-1. Full text in Research Archive Show summary

  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.

• Spedener, Mélanie; Tofastrud, Morten; Devineau, Olivier & Zimmermann, Barbara (2019). Microhabitat selection of free-ranging beef cattle in south-boreal forest. Applied Animal Behaviour Science. ISSN 0168-1591. 213, p. 33–39. doi: 10.1016/j.applanim.2019.02.006. Full text in Research Archive
• Fuchs, Boris; Sørheim, Kristin; Chincarini, Matteo; Brunberg, Emma; Stubsjøen, Solveig Marie & Bratbergsengen, Kjell [Show all 10 contributors for this article] (2019). Heart rate sensor validation and seasonal and diurnal variation of body temperature and heart rate in domestic sheep. Veterinary and Animal Science. 8, p. 1–9. doi: 10.1016/j.vas.2019.100075. Full text in Research Archive Show summary

  Advantages of low input livestock production on large pastures, including animal welfare, biodiversity and low production costs are challenged by losses due to undetected disease, accidents and predation. Precision livestock farming (PLF) enables remote monitoring on individual level with potential for predictive warning. Body temperature (Tb) and heart rate (HR) could be used for early detection of diseases, stress or death. We tested physiological sensors in free-grazing Norwegian white sheep in Norway. Forty Tb sensors and thirty HR sensors were surgically implanted in 40 lambs and 10 ewes. Eight (27%) of the HR and eight (20%) of the Tb sensors were lost during the study period. Two Tb sensors migrated from the abdominal cavity in to the digestive system. ECG based validation of the HR sensors revealed a measurement error of 0.2 bpm (SD 5.2 bpm) and correct measurement quality was assigned in 90% of the measurements. Maximum and minimum HR confirmed by ECG was 197 bpm and 68 bpm respectively. Mean passive HR was 90 bpm (SD=13 bpm) for ewes and 112 bpm (SD=13 bpm) for lambs. Mean Tb for all animals was 39.6°C (range 36.9 to 41.8°C). Tb displayed 24-hour circadian rhythms during 80.7 % but HR only during 41.0 % of the studied period. We established baseline values and conclude that these sensors deliver good quality. For a wide agricultural use, the sensor implantation method has to be further developed and real-time communication technology added.

• Tofastrud, Morten; Devineau, Olivier & Zimmermann, Barbara (2019). Habitat selection of free-ranging cattle in productive coniferous forests of south-eastern Norway. Forest Ecology and Management. ISSN 0378-1127. 437, p. 1–9. doi: 10.1016/j.foreco.2019.01.014. Full text in Research Archive
• Sebastián-González, Esther; Barbosa, Jomar Magalhães; Pérez-García, Juan M.; Morales-Reyes, Zebensui; Botella, Francisco & Olea, Pedro P. [Show all 45 contributors for this article] (2019). Scavenging in the Anthropocene: Human impact drives vertebrate scavenger species richness at a global scale. Global Change Biology. ISSN 1354-1013. 25(9), p. 3005–3017. doi: 10.1111/gcb.14708. Full text in Research Archive
• Narvaez-Ortiz, Heidy Y.; Moreno, Andrea Johana Lopez; Gupta, Nishith & Zimmermann, Barbara (2018). A CTP synthase undergoing stage-specific spatial expression is essential for the survival of the intracellular parasite Toxoplasma gondii. Frontiers in Cellular and Infection Microbiology. 8:83, p. 1–17. doi: 10.3389/fcimb.2018.00083. Full text in Research Archive
• Breisjøberget, Jo Inge; Odden, Morten; Wegge, Per; Zimmermann, Barbara & Andreassen, Harry Peter (2018). The alternative prey hypothesis revisited: Still valid for willow ptarmigan population dynamics. PLOS ONE. 13(6). doi: 10.1371/journal.pone.0197289. Full text in Research Archive
• Milleret, Cyril Pierre; Fernandez, Andres Avelino Ordiz; Chapron, Guillaume; Andreassen, Harry Peter; Kindberg, Jonas & Månsson, Johan [Show all 12 contributors for this article] (2018). Habitat segregation between brown bears and grey wolves in a human-dominated landscape. Ecology and Evolution. 8(23), p. 11450–11466. doi: 10.1002/ece3.4572. Full text in Research Archive
• Meisingset, Erling L.; Loe, Leif Egil; Brekkum, Øystein; Bischof, Richard; Rivrud, Inger Maren & Lande, Unni Støbet [Show all 9 contributors for this article] (2018). Spatial mismatch between management units and movement ecology of a partially migratory ungulate. Journal of Applied Ecology. ISSN 0021-8901. 55(2), p. 745–753. doi: 10.1111/1365-2664.13003. Full text in Research Archive Show summary

  1. Population-level management is difficult to achieve if wildlife routinely crosses administrative boundaries, as is particularly frequent for migratory populations. However, the degree of mismatch between management units and scales at which ecological processes operate has rarely been quantified. Such insight is vital for delimiting functional population units of partially migratory species common in northern forest ecosystems. 2. We combined an extensive dataset of 412 GPS-marked red deer (Cervus elaphus) across Norway with information on the size and borders of two administrative levels, the governmental level (municipality) and landowner level (local management units, LMUs), to determine the timing and scale of mismatch between animal space use and management units. We analysed how landscape characteristics affected the use of management units and the timing and likelihood of crossing borders between them, in an effort to delineate more appropriate units in various landscapes. 3. Median municipality size could potentially cover 70% of female and 62% of male annual ranges, while only 12% and 4% of LMUs were expansive enough to accommodate migratory routes in females and males, respectively. Red deer migrate along elevational gradients and are more likely to find both suitable lowland winter habitat and higher summer habitat within management units with variable topography. Consistent with this, the likelihood of border crossing decreased with increasing diversity of elevations. 4. Synthesis and applications. We demonstrate a considerable mismatch between animal space use and management units. Far-ranging movements and frequent administrative border crossings during autumn migration coincides with the period of active management (hunting season). Our study also highlights that, due to extensive movements of males, coordination of management aims may provide a more realistic avenue than increasing sizes of local management units. A more general insight is that the degree of mismatch between range use and management units depends on the season and landscape type. This needs to be accounted for when delimitating functional population units of migratory populations.

• Tofastrud, Morten; Hegnes, Hilde; Devineau, Olivier & Zimmermann, Barbara (2018). Activity patterns of free-ranging beef cattle in Norway. Acta agriculturæ Scandinavica. Section A, Animal science. ISSN 0906-4702. p. 1–9. doi: 10.1080/09064702.2018.1524928. Full text in Research Archive
• Eriksen, Ane; Wabakken, Petter; Maartmann, Erling & Zimmermann, Barbara (2018). Den site selection by male brown bears at the population’s expansion front. PLOS ONE. 13(8), p. 1–18. doi: 10.1371/journal.pone.0202653. Full text in Research Archive
• Recio, Mariano R; Zimmermann, Barbara; Wikenros, Camilla; Zetterberg, Andreas; Wabakken, Petter & Sand, Håkan (2018). Data to model risks for recolonizing wolves in Scandinavia through the integration of territory presence and human-driven mortalities. Data in Brief. 20, p. 686–690. doi: 10.1016/j.dib.2018.08.060. Full text in Research Archive
• Perez, Ana Sanz; Fernandez, Andres Avelino Ordiz; Sand, Håkan; Swenson, Jon; Wabakken, Petter & Wikenros, Camilla [Show all 9 contributors for this article] (2018). No place like home? A test of the natal habitat-biased dispersal hypothesis in Scandinavian wolves. Royal Society Open Science. 5(12). doi: 10.1098/rsos.181379. Full text in Research Archive Show summary

  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.

• Recio, Mariano R; Zimmermann, Barbara; Wikenros, Camilla; Zetterberg, Andreas; Wabakken, Petter & Sand, Håkan (2018). Integrated spatially-explicit models predict pervasive risks to recolonizing wolves in Scandinavia from human-driven mortality. Biological Conservation. ISSN 0006-3207. 226, p. 111–119. doi: 10.1016/j.biocon.2018.07.025. Full text in Research Archive
• Chapron, Guillaume; Wikenros, Camilla; Liberg, Olof; Wabakken, Petter; Flagstad, Øystein & Milleret, Cyril [Show all 11 contributors for this article] (2016). Estimating wolf (Canis lupus) population size from number of packs and an individual based model. Ecological Modelling. ISSN 0304-3800. 339, p. 33–44. doi: 10.1016/j.ecolmodel.2016.08.012. Full text in Research Archive
• Rivrud, Inger Maren; Bischof, Richard; Meisingset, Erling L.; Zimmermann, Barbara; Loe, Leif Egil & Mysterud, Atle (2016). Leave before it's too late: Anthropogenic and environmental triggers of autumn migration in a hunted ungulate population. Ecology. ISSN 0012-9658. 97(4), p. 1058–1068. doi: 10.1890/15-1191.1. Full text in Research Archive
• Sand, Håkan; Eklund, Ann; Zimmermann, Barbara; Wikenros, Camilla & Wabakken, Petter (2016). Prey Selection of Scandinavian Wolves: Single Large or Several Small? PLOS ONE. 11(12). doi: 10.1371/journal.pone.0168062. Full text in Research Archive
• Fuchs, Boris; Zimmermann, Barbara; Wabakken, Petter; Bornstein, Set; Månsson, Johan & Evans, Alina [Show all 11 contributors for this article] (2016). Sarcoptic mange in the Scandinavian wolf Canis lupus population. BMC Veterinary Research. 12(156). doi: 10.1186/s12917-016-0780-y. Full text in Research Archive Show summary

  Background: Sarcoptic mange, a parasitic disease caused by the mite Sarcoptes scabiei, is regularly reported on wolves Canis lupus in Scandinavia. We describe the distribution and transmission of this parasite within the small but recovering wolf population by analysing 269 necropsy reports and performing a serological survey on 198 serum samples collected from free-ranging wolves between 1998 and 2013. Results: The serological survey among 145 individual captured Scandinavian wolves (53 recaptures) shows a consistent presence of antibodies against sarcoptic mange. Seropositivity among all captured wolves was 10.1 % (CI. 6.4 %–15.1 %). Sarcoptic mange-related mortality reported at necropsy was 5.6 % and due to secondary causes, predominantly starvation. In the southern range of the population, seroprevalence was higher, consistent with higher red fox densities. Female wolves had a lower probability of being seropositive than males, but for both sexes the probability increased with pack size. Recaptured individuals changing from seropositive to seronegative suggest recovery from sarcoptic mange. The lack of seropositive pups (8–10 months, N = 56) and the occurrence of seropositive and seronegative individuals in the same pack indicates interspecific transmission of S. scabiei into this wolf population. Conclusions: We consider sarcoptic mange to have little effect on the recovery of the Scandinavian wolf population. Heterogenic infection patterns on the pack level in combination with the importance of individualbased factors (sex, pack size) and the north–south gradient for seroprevalence suggests low probability of wolf-to-wolf transmission of S. scabiei in Scandinavia.

• Mayer, Martin; Brown, Gregory P.; Zimmermann, Barbara & Shine, Richard (2015). High infection intensities, but negligible fitness costs, suggest tolerance of gastrointestinal nematodes in a tropical snake. Austral ecology (Print). ISSN 1442-9985. 40(6), p. 683–692. doi: 10.1111/aec.12235. Full text in Research Archive Show summary

  We investigated patterns of prevalence and intensity of gastrointestinal nematode infections in a tropical natricine snake, the keelback (Tropidonophis mairii). Ninety-eight per cent of keelbacks were infected with Tanqua anomala (Gnathostomidae), with infection intensities of up to 243 worms per snake. Infection with T. anomala caused severe inflammation of stomach mucosa and submucosa at the sites of parasite attachment and encystment. Nonetheless, we did not detect detrimental effects of nematode infection on measures of fitness among wild or captive snakes. Snakes with heavier nematode infections had higher body condition scores than less-infected individuals. Deworming captive snakes had no measurable effect on their growth rate, body condition or locomotor performance. In combination with an earlier study on blood-dwelling hepatozoons, our work suggests that keelbacks have a high tolerance to parasites. The ‘fast-pace’ life history and short lifespan of these snakes may make it beneficial for them to tolerate infection, rather than expend energy on resisting parasite attack.

• Mayer, Martin; Brown, Gregory P.; Zimmermann, Barbara; Greenlees, Matthew J. & Shine, Richard (2015). Habitat use of the introduced cane toad (Rhinella marina) and native frog species in tropical Australia. Journal of Tropical Ecology. ISSN 0266-4674. 31(6), p. 531–540. doi: 10.1017/S0266467415000474. Full text in Research Archive Show summary

  The ecological impacts of introduced species can reveal mechanisms underlying habitat selection and behaviour. We investigated the habitat use of native frog species and the invasive cane toads (Rhinella marina) in tropical northern Australia to measure overlap in habitat use, and to test if the presence of the cane toad influences frog behaviour. Native frog species and the cane toad both preferred habitats close to water and unvegetated holes. However, native frogs were found further from water (on average 19.4 m) than were toads (on average 12.6 m), and preferred areas with higher vegetation (8–50 cm) than did toads,which were more abundant in vegetation lower than 8 cm. For both types of anuran, the next neighbour was more often of the same type (89% in frogs, 52% in toads) than expected by chance (observed ratio: 75% frogs vs 25% toads), reflecting these differences in habitat use. Our Counts of frog abundance increased on average 14.5% in areas from which we removed cane toads temporarily. This result suggests that cane toads inhibit the activity of native anurans either by inducing avoidance, or by reducing activity. By modifying the behaviour and spatial distribution of native taxa, invasive cane toads may curtail activities such as feeding and breeding.

• Zimmermann, Barbara; Sand, Håkan; Wabakken, Petter; Liberg, Olof & Andreassen, Harry Peter (2015). Predator‐dependent functional response in wolves: from food limitation to surplus killing. Journal of Animal Ecology. ISSN 0021-8790. 84(1), p. 102–112. doi: 10.1111/1365-2656.12280. Full text in Research Archive Show summary

  1. The functional response of a predator describes the change in per capita kill rate to changes in prey density. This response can be influenced by predator densities, giving a predator- dependent functional response. In social carnivores which defend a territory, kill rates also depend on the individual energetic requirements of group members and their contribution to the kill rate. 2. This study aims to provide empirical data for the functional response of wolves Canis lupus to the highly managed moose Alces alces population in Scandinavia. We explored prey and predator dependence, and how the functional response relates to the energetic requirements of wolf packs. 3. Winter kill rates of GPS-collared wolves and densities of cervids were estimated for a total of 22 study periods in 15 wolf territories. The adult wolves were identified as the individuals responsible for providing kills to the wolf pack, while pups could be described as inept hunters. 4. The predator-dependent, asymptotic functional response models (i.e. Hassell–Varley type II and Crowley–Martin) performed best among a set of 23 competing linear, asymptotic and sigmoid models. Small wolf packs acquired >3 times as much moose biomass as required to sustain their field metabolic rate (FMR), even at relatively low moose abundances. Large packs (6–9 wolves) acquired less biomass than required in territories with low moose abundance. 5. We suggest the surplus killing by small packs is a result of an optimal foraging strategy to consume only the most nutritious parts of easy accessible prey while avoiding the risk of being detected by humans. Food limitation may have a stabilizing effect on pack size in wolves, as supported by the observed negative relationship between body weight of pups and pack size.

• Sente, Celsus; Meisingset, Erling L.; Evans, Alina; Wedul, Sari Jemiina; Zimmermann, Barbara & Arnemo, Jon Martin (2014). Reversible Immobilization of Free-ranging Red Deer (Cervus elaphus) with Xylazine-Tiletamine-Zolazepam and Atipamezole. Journal of Wildlife Diseases. ISSN 0090-3558. 50(2), p. 359–363. doi: 10.7589/2012-10-267. Full text in Research Archive
• Zimmermann, Barbara; Nelson, Lindsey; Wabakken, Petter; Sand, Håkan & Liberg, Olof (2014). Behavioral responses of wolves to roads: scale-dependent ambivalence. Behavioral Ecology. ISSN 1045-2249. 25(6), p. 1353–1364. doi: 10.1093/beheco/aru134. Full text in Research Archive Show summary

  Throughout their recent recovery in several industrialized countries, large carnivores have had to cope with a changed landscape dominated by human infrastructure. Population growth depends on the ability of individuals to adapt to these changes by making use of new habitat features and at the same time to avoid increased risks of mortality associated with human infrastructure. We analyzed the summer movements of 19 GPS-collared resident wolves (Canis lupus L.) from 14 territories in Scandinavia in relation to roads. We used resource and step selection functions, including >12 000 field-checked GPS-positions and 315 kill sites. Wolves displayed ambivalent responses to roads depending on the spatial scale, road type, time of day, behavioral state, and reproductive status. At the site scale (approximately 0.1 km2), they selected for roads when traveling, nearly doubling their travel speed. Breeding wolves moved the fastest. At the patch scale (10 km2), house density rather than road density was a significant negative predictor of wolf patch selection. At the home range scale (approximately 1000 km2), breeding wolves increased gravel road use with increasing road availability, although at a lower rate than expected. Wolves have adapted to use roads for ease of travel, but at the same time developed a cryptic behavior to avoid human encounters. This behavioral plasticity may have been important in allowing the successful recovery of wolf populations in industrialized countries. However, we emphasize the role of roads as a potential cause of increased human-caused mortality.

• Zimmermann, Barbara; Wedul, Sari Jemiina; Johnsen, Kaja; Strømseth, Thomas Holm & Østerhus, Benedicte (2014). Hjortens trekkmønster skaper forvaltningsutfordringer i Hedmark. In Storaas, Torstein & Langdal, Kjell (Ed.), Ikkje berre ulv og bly - glimt frå forskninga på Evenstad. Oplandske Bokforlag. ISSN 9788275182232. p. 145–159. Full text in Research Archive Show summary

  Jaktstatistikken fra Hedmark fylke tilsier at hjorten er tilbake for godt i Hedmark og befinner seg i en sterk vekstfase. Med det følger mange forvaltningsutfordringer. Vi har studert hjortens forflytningsmønster og trekkatferd ved å merke 51 hinder med GPS på fire merkeplasser i henholdsvis Rendalen, Stor-Elvdal, Åmot og Stange kommune. De aller fleste hindene oppholdt seg i et sommerområde som var atskilt fra vinterområdet. Avstanden mellom de to områdene var gjennomsnittlig lenger enn for hinder merket på Vest- og Sørlandet. For tre av hindene var avstanden rundt 100 km. På trekkruten var hindene innom forskjellige kommuner, og totalt ble vårt studieområde utvidet fra de fire vinterkommunene til 29 kommuner for hele året. Vi anbefaler derfor en mer regional forvaltning av hjorten enn dagens forvaltning på eiendoms- eller kommunenivå. Innmark og fôringsplasser ble flittig brukt av hindene, og da spesielt nattestid. Overvåkning av fôringsplasser for elg som opprinnelig ble etablert for å redusere trafikkulykker og beiteskader i skogen, viste at hjorten var til stede på samtlige plasser langs Glommas hoveddalføre i Stor-Elvdal og Rendalen kommuner. Mye tyder på at hjorten har tatt over flere plasser, og at elg og hjort til en viss grad unngikk hverandre i tid og rom. Det kan tenkes at hjorten kan minske den ønskede effekten av vinterfôring av elg. Hvis vinterfôring skal fortsette, anbefaler vi å legge ut flere siloballer samtidig per fôringsplass for å sikre elgens tilgang til fôr.

• Zimmermann, Barbara; Sand, Håkan; Wabakken, Petter; Wikenros, Camilla; Eriksen, Ane & Strømseth, Thomas Holm [Show all 13 contributors for this article] (2014). Ulven som rovdyr på klauvvilt i Skandinavia. In Storaas, Torstein & Langdal, Kjell (Ed.), Ikkje berre ulv og bly - glimt frå forskninga på Evenstad. Oplandske Bokforlag. ISSN 9788275182232. p. 221–245. Full text in Research Archive Show summary

  Med ulvens tilbakekomst og bestandsøkning i Skandinavia spurte mange om ulven ville påvirke klauvviltbestanden og redusere elgjaktens avkastning. Det Skandinaviske Ulveforskningsprosjektet SKANDULV (http://skandulv.nina.no/) har derfor fokusert mye av forskningen på ulvens rolle som rovdyr. Ved å merke ulv med GPS og lete etter byttedyrrester på GPS-posisjonene kunne vi konstatere at elg var det viktigste byttedyret til ulv, og at rådyr bare hadde betydning i områder med svært høye rådyrtettheter. Blant elgene selekterte ulv for kalver og de eldste dyrene i bestanden. Ofte var de gamle byttedyrene i dårlig hold. Drapsraten lå på gjennomsnittlig 0,25 (vinter) og 0,58 (sommer) elg per flokk per døgn og varierte med tilgang på elg og flokkstørrelse. Ulvepar uten valper og små ulveflokker la ned mer bytte enn nødvendig for å dekke sine daglige energibehov, mens i store flokker klarte ikke ulveparet å legge ned nok bytte til alle flokkmedlemmene. Totalt vil ulveflokker ta 100 – 144 elger i løpet av et år. Hvor stor del av elgene i et jaktområde som blir drept av ulv, avhenger av ulverevirets størrelse og elgens tetthet i området. SKANDULV har utviklet et web-basert verktøy for rettighetshavere for å forutsi jaktens avkastning i områder med ulv og/eller bjørn ved forskjellige høstingsregimer (http://www.algforvaltning.se/moosemodel/). Denne nye kunnskapen om ulvenes atferd og påvirkning på elgbestanden i Skandinavia er viktig for forvaltningen. I denne artikkelen belyser vi flere aspekter ved ulvens rolle som rovdyr i det kompliserte samspillet med landskapet, flere rovdyrarter, åtselsetere, klauvvilt og mennesker.

• Zimmermann, Barbara (2013). How does animal monitoring with GPS tags contribute to ecology and conservation? Tidsskriftet Utmark. Full text in Research Archive
• Gervasi, Vincenzo; Sand, Håkon; Zimmermann, Barbara; Mattisson, Jenny; Wabakken, Petter & Linnell, John Durrus (2013). Decomposing risk: Landscape structure and wolf behavior generate different predation patterns in two sympatric ungulates. Ecological Applications. ISSN 1051-0761. 23(7), p. 1722–1734. doi: 10.1890/12-1615.1. Full text in Research Archive Show summary

  Recolonizing carnivores can have a large impact on the status of wild ungulates, which have often modified their behavior in the absence of predation. Therefore, understanding the dynamics of reestablished predator–prey systems is crucial to predict their potential ecosystem effects. We decomposed the spatial structure of predation by recolonizing wolves (Canis lupus) on two sympatric ungulates, moose (Alces alces) and roe deer (Capreolus capreolus), in Scandinavia during a 10-year study. We monitored 18 wolves with GPS collars, distributed over 12 territories, and collected records from predation events. By using conditional logistic regression, we assessed the contributions of three main factors, the utilization patterns of each wolf territory, the spatial distribution of both prey species, and fine-scale landscape structure, in determining the spatial structure of moose and roe deer predation risk. The reestablished predator–prey system showed a remarkable spatial variation in kill occurrence at the intra-territorial level, with kill probabilities varying by several orders of magnitude inside the same territory. Variation in predation risk was evident also when a spatially homogeneous probability for a wolf to encounter a prey was simulated. Even inside the same territory, with the same landscape structure, and when exposed to predation by the same wolves, the two prey species experienced an opposite spatial distribution of predation risk. In particular, increased predation risk for moose was associated with open areas, especially clearcuts and young forest stands, whereas risk was lowered for roe deer in the same habitat types. Thus, fine-scale landscape structure can generate contrasting predation risk patterns in sympatric ungulates, so that they can experience large differences in the spatial distribution of risk and refuge areas when exposed to predation by a recolonizing predator. Territories with an earlier recolonization were not associated with a lower hunting success for wolves. Such constant efficiency in wolf predation during the recolonization process is in line with previous findings about the naı¨ve nature of Scandinavian moose to wolf predation. This, together with the human-dominated nature of the Scandinavian ecosystem, seems to limit the possibility for wolves to have large ecosystem effects and to establish a behaviorally mediated trophic cascade in Scandinavia Alces alces, Canis lupus, Capreolus capreolus, catchability, dilution effect, kill occurrence, moose, predation risk, predator recolonization, prey refugia, roe deer, Scandinavia

• Nilsen, Erlend Birkeland; Christianson, David; Gaillard, Jean-Michel; Halley, Duncan John; Linnell, John Durrus & Odden, Morten [Show all 9 contributors for this article] (2012). Describing food habits and predation: field methods and statistical considerations. In Boitani, Luigi & Powell, Roger A. (Ed.), Carnivore ecology and conservation. A handbook of techniques. Oxford University Press. ISSN 9780199558537. p. 256–272. doi: 10.1093/oso/9780199558520.003.0011. Full text in Research Archive
• Gervasi, Vincenzo; Nilsen, Erlend Birkeland; Sand, Håkan; Panzacchi, Manuela; Rauset, Geir Rune & Pedersen, Hans-Christian [Show all 13 contributors for this article] (2012). Predicting the potential demographic impact of predators on their prey: a comparative analysis of two carnivore-ungulate systems in Scandinavia. Journal of Animal Ecology. ISSN 0021-8790. 81(2), p. 443–454. doi: 10.1111/j.1365-2656.2011.01928.x. Full text in Research Archive
• Sand, Håkan; Vucetich, John A.; Zimmermann, Barbara; Wabakken, Petter; Wikenros, Camilla & Pedersen, Hans-Christian [Show all 8 contributors for this article] (2012). Assessing the influence of prey-predator ratio, prey age structure and packs size on wolf kill rates. Oikos. ISSN 0030-1299. 121(9), p. 1454–1463. doi: 10.1111/j.1600-0706.2012.20082.x. Full text in Research Archive
• Bischof, Richard; Loe, Leif Egil; Meisingset, Erling; Zimmermann, Barbara; Moorter, Bram F. A. Van & Mysterud, Atle (2012). A Migratory Northern Ungulate in the Pursuit of Spring: Jumping or Surfing the Green Wave? The American Naturalist. ISSN 0003-0147. 180(4), p. 407–424. doi: 10.1086/667590. Full text in Research Archive Show summary

  The forage-maturation hypothesis (FMH) states that herbivores migrate along a phenological gradient of plant development in order to maximize energy intake. Despite strong support for the FMH, the actual relationship between plant phenology and ungulate movement has remained enigmatic. We linked plant phenology (MODIS–normalized difference vegetation index [NDVI] data) and space use of 167 migratory and 78 resident red deer (Cervus elaphus), using a space-time-time matrix of “springness,” defined as the instantaneous rate of green-up. Consistent with the FMH, migrants experienced substantially greater access to early plant phenology than did residents. Deer were also more likely to migrate in areas where migration led to greater gains in springness. Rather than “surfing the green wave” during migration, migratory red deer moved rapidly from the winter to the summer range, thereby “jumping the green wave.” However, migrants and, to a lesser degree, residents did track phenological green-up through parts of the growing season by making smaller-scale adjustments in habitat use. Despite pronounced differences in their life histories, we found only marginal differences between male and female red deer in this study. Those differences that we did detect pointed toward additional constraints on female space-use tactics, such as those posed by calving and caring for dependent offspring. We conclude that whereas in some systems migration itself is a way to surf the green wave, in others it may simply be a means to reconnect with phenological spring at the summer range. In the light of ubiquitous anthropogenic environmental change, understanding the relationship between the green wave and ungulate space use has important consequences for the management and conservation of migratory ungulates and the phenomenon of migration itself. © 2012 by The University of Chicago.

• Mysterud, Atle; Loe, Leif Egil; Zimmermann, Barbara; Bischof, Richard; Veiberg, Vebjørn & Meisingset, Erling (2011). Partial migration in expanding red deer populations at northern latitudes – a role for density dependence? Oikos. ISSN 0030-1299. 120(12), p. 1817–1825. doi: 10.1111/j.1600-0706.2011.19439.x. Full text in Research Archive
• Torkar, Gregor; Zimmermann, Barbara & Willebrand, Tomas (2011). Qualitative interviews in human dimensions studies about nature conservation. Varstvo Narave. ISSN 0506-4252. 25, p. 39–52. Full text in Research Archive
• Eriksen, Ane; Wabakken, Petter; Zimmermann, Barbara; Andreassen, Harry Peter; Arnemo, Jon Martin & Gundersen, Hege [Show all 13 contributors for this article] (2011). Activity patterns of predator and prey: a simultaneous study of GPS-collared wolves and moose. Animal Behaviour. ISSN 0003-3472. 81(2), p. 423–431. doi: 10.1016/j.anbehav.2010.11.011. Full text in Research Archive Show summary

  We studied the simultaneous activity patterns of a breeding wolf, Canis lupus, pair and five adult moose, Alces alces, cows from April to November 2004 in a wolf territory in southeastern Norway. All study animals were GPS collared, and we used a total of 8297 fixes to analyse their temporal activity patterns. We examined the daily activity rhythm of the two species and how this varied seasonally through the study period, and investigated the association in activity patterns between the two species. Wolf activity peaked at dawn. The distance moved per time unit and the linearity of the movement were higher in wolves than in moose, but both species showed a decrease in these variables in June, coinciding with the denning and calving seasons. With the exception of the summer months when the activity of wolves and moose was limited by the raising of offspring, we found no correlation between the temporal activities of the two species. Hence, we did not find support for the hypothesis that the wolves and moose in Scandinavia have adjusted to each other’s activity patterns. We discuss the results in light of the relative density of the two species and the intensive human harvest of moose, and hypothesize that synchronicity of predator–prey activity patterns may be ratio dependent.

• Merrill, Evelyn; Sand, Håkan; Zimmermann, Barbara; McPhee, Heather; Webb, Nathan & Hebblewhite, Mark [Show all 8 contributors for this article] (2010). Building a mechanistic understanding of predation with GPS-based movement data. Philosophical Transactions of the Royal Society of London. Biological Sciences. ISSN 0962-8436. 365(1550), p. 2279–2288. doi: 10.1098/rstb.2010.0077. Full text in Research Archive Show summary

  Quantifying kill rates and sources of variation in kill rates remains an important challenge in linking predators to their prey. We address current approaches to using global positioning system (GPS)-based movement data for quantifying key predation components of large carnivores. We review approaches to identify kill sites from GPS movement data as a means to estimate kill rates and address advantages of using GPS-based data over past approaches. Despite considerable progress, modelling the probability that a cluster of GPS points is a kill site is no substitute for field visits, but can guide our field efforts. Once kill sites are identified, time spent at a kill site (handling time) and time between kills (killing time) can be determined. We show how statistical models can be used to investigate the influence of factors such as animal characteristics (e. g. age, sex, group size) and landscape features on either handling time or killing efficiency. If we know the prey densities along paths to a kill, we can quantify the `attack success' parameter in functional response models directly. Problems remain in incorporating the behavioural complexity derived from GPS movement paths into functional response models, particularly in multi-prey systems, but we believe that exploring the details of GPS movement data has put us on the right path.

• Eriksen, Ane; Wabakken, Petter; Zimmermann, Barbara; Andreassen, Harry Peter; Arnemo, Jon Martin & Gundersen, Hege [Show all 13 contributors for this article] (2009). Encounter frequencies between GPS-collared wolves (Canis lupus) and moose (Alces alces) in a Scandinavian wolf territory. Ecological Research. ISSN 0912-3814. 24(3), p. 547–557. doi: 10.1007/s11284-008-0525-x. Full text in Research Archive
• Gundersen, Hege; Solberg, Erling Johan; Wabakken, Petter; Storaas, Torstein; Zimmermann, Barbara & Andreassen, Harry Peter (2008). Three approaches to estimate wolf Canis lupus predation rates on moose Alces alces populations. European Journal of Wildlife Research. ISSN 1612-4642. 54. doi: 10.1007/s10344-007-0155-6. Full text in Research Archive Show summary

  We employed three different methods to estimate predation rates on moose in a newly colonized wolf territory in Norway. In the first two methods we estimated predation rates based on the difference in calf:cow ratios outside and inside the wolf pack territory from 1) hunter observations and 2) aerial surveys. In the last method (3) we estimated loss of calves of radio-collared cows inside and outside the wolf pack territory. The difference in mortality rates estimated between the area subject to predation and the area outside the wolf pack territory essentially constitutes the additive component of predation. We also tested the sensitivity of violating the assumptions of method 1 and 2 related to equal fecundity and mortality due to other factors than predation inside and outside the wolf pack territory. Predation rates varied considerably between years and methods used, with hunter observations (method 1) giving the lowest and aerial surveys (method 2) giving the highest estimates. Method 3 (radio telemetry) was the most direct assessment of predation and probably the best approach to estimate predation rates in moose. However, all three methods show the same yearly changes and may therefore be appropriate to question trends trough time or between areas.

• Sand, Håkan; Wabakken, Petter; Zimmermann, Barbara; Johansson, Örjan; Pedersen, Hans Christian & Liberg, Olof (2008). Summer kill rates and predation pattern in a wolf-moose system: can we rely on winter estimates? Oecologia. ISSN 0029-8549. 156(1), p. 53–64. Full text in Research Archive Show summary

  So far the vast majority of studies on large carnivore predation, including kill rates and consumption, have been based on winter studies. Because large carnivores relying on ungulates as prey often show a preference for juveniles, kill rates may be both higher and more variable during the summer season than during the rest of the year leading to serious underestimates of the total annual predation rate. This study is the first to present detailed empirical data on kill rates and prey selection in a wolf-moose system during summer (June-September) as obtained by applying modern Global Positioning System-collar techniques on individual wolves (Canis lupus) in Scandinavia. Moose (Alces alces) was the dominant prey species both by number (74.4%) and biomass (95.6%); 89.9% of all moose killed were juveniles, representing 76.0% of the biomass consumed by wolves. Kill rate in terms of the kilogram biomass/kilogram wolf per day averaged 0.20 (range: 0.07-0.32) among wolf territories and was above, or well above, the daily minimum food requirements in most territories. The average number of days between moose kills across wolf territories and study periods was 1.71 days, but increased with time and size of growing moose calves during summer. Over the entire summer (June-September, 122 days), a group (from two to nine) of wolves killed a total of 66 (confidence interval 95%; 56-81) moose. Incorporation of body growth functions of moose calves and yearlings and wolf pups over the summer period showed that wolves adjusted their kill rate on moose, so the amount of biomass/kilogram wolf was relatively constant or increased. The kill rate was much higher (94-116%) than estimated from the winter period. As a consequence, projecting winter kill rates to obtain annual estimates of predation in similar predator-prey systems may result in a significant underestimation of the total number of prey killed.

• May, Roel; Dijk, Jiska Joanneke van; Wabakken, Petter; Swenson, Jon; Linnell, John D.C. & Zimmermann, Barbara [Show all 10 contributors for this article] (2008). Habitat differentiation within the large-carnivore community of Norway's multiple-use landscapes. Journal of Applied Ecology. ISSN 0021-8901. 45(5), p. 1382–1391. doi: 10.1111/j.1365-2664.2008.01527.x. Full text in Research Archive
• Wabakken, Petter; Sand, Håkan; Kojola, Ilpo; Zimmermann, Barbara; Arnemo, Jon Martin & Pedersen, Hans Christian [Show all 7 contributors for this article] (2007). Multistage, long-range natal dispersal by a Global Positioning System-collared Scandinavian wolf. Journal of Wildlife Management. ISSN 0022-541X. 71(5), p. 1631–1634. Full text in Research Archive Show summary

  We document a new record dispersal for wolves worldwide. The natal straight-line dispersal distance of a Global Positioning System-collared female wolf from the Scandinavian population was 1,092 km from southeast Norway to northeast Finland, with a multistage actual travel distance of > 10,000 km. Natural gene flow to the isolated, inbred Scandinavian wolf population may occur if survival of dispersers is improved.

• Zimmermann, Barbara; Wabakken, Petter; Sand, Håkan; Pedersen, Hans Christian & Liberg, Olof (2007). Wolf movement patterns: a key to estimation of kill rate? Journal of Wildlife Management. ISSN 0022-541X. 71(4), p. 1177–1182. Full text in Research Archive Show summary

  To estimate wolf (Canis lupus) kill rates from fine-scale movement patterns, we followed adult wolves in 3 territories of the Scandinavian wolf population using Global Positioning Systems (GPS) during the winters of 2001-2003. The resulting 6 datasets of 62-84 study days gave a total of 8,747 hourly GPS positions. We visited clusters of positions in the field on average 8.8 days after positioning and found moose (Alces alces) killed by wolves during the study period on 74 (8%) of the 953 dusters. The number of positions and visits to a cluster, their interaction, and the proportion of afternoon positions were significant fixed effects in mixed logistic-regression models predicting the probability of a cluster containing a wolf-killed moose. The models, however, displayed a poor goodness-of-fit and were not a suitable tool for estimating kill rates from positioning data alone. They might be used to reduce fieldwork by excluding unlikely clusters, although the reduction was not substantial. We discuss proximate factors (i.e., human disturbance and access to prey) as well as ultimate factors (i.e., social organization, intra-guild dominance, and litter size) as potential causes of the observed high temporal and spatial variation in prey-handling. For similar future kill-rate studies, we recommend increasing field efforts and shortening positioning intervals.

• Gundersen, Hege; Solberg, Erling Johan; Wabakken, Petter; Storaas, Torstein; Zimmermann, Barbara & Andreassen, Harry P. (2007). Three approaches to estimate wolf Canis lupus predation rates on moose Alces alces populations. European Journal of Wildlife Research. ISSN 1612-4642. doi: 10.1007/s10344-007-0155-6. Full text in Research Archive Show summary

  We employed three different methods to estimate predation rates on moose in a newly colonized wolf territory in Norway. In the first two methods we estimated predation rates based on the difference in calf:cow ratios outside and inside the wolf pack territory from 1) hunter observations and 2) aerial surveys. In the last method (3) we estimated loss of calves of radio-collared cows inside and outside the wolf pack territory. The difference in mortality rates estimated between the area subject to predation and the area outside the wolf pack territory essentially constitutes the additive component of predation. We also tested the sensitivity of violating the assumptions of method 1 and 2 related to equal fecundity and mortality due to other factors than predation inside and outside the wolf pack territory. Predation rates varied considerably between years and methods used, with hunter observations (method 1) giving the lowest and aerial surveys (method 2) giving the highest estimates. Method 3 (radio telemetry) was the most direct assessment of predation and probably the best approach to estimate predation rates in moose. However, all three methods show the same yearly changes and may therefore be appropriate to question trends trough time or between areas. Original Paper: Accepted 29 October 2007 Published online: 27 November 2007

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• Panzacchi, Manuela; Moorter, Bram Van; Stange, Erik; Simensen, Trond; Sydenham, Markus A. K. & Thorsen, Neri Horntvedt [Show all 13 contributors for this article] (2026). GreenPlan - Lansering av ny kartportal for Grønn Infrastruktur i Norge og samutvikling av støtteverktøy for arealprioritering, scenario- og konsekvensvurderinger. DEL1 - STATUS HJORT. Full text in Research Archive
• Zimmermann, Barbara (2025). Ett år med elgku E2102. Full text in Research Archive
• Zimmermann, Barbara (2025). The life in boxes and bubbles: Research groups. Full text in Research Archive
• Zimmermann, Barbara (2025). Elgjakt, rovdyr og skogbruk. Full text in Research Archive
• Zimmermann, Barbara (2025). Elg i grenseland. Full text in Research Archive
• Zimmermann, Barbara (2025). Nyeste SKANDULV-forskning på ulv. Full text in Research Archive
• Zimmermann, Barbara (2025). Evenstads viltforskning og IWC 2025. Full text in Research Archive
• Zimmermann, Barbara (2025). Betydningen av lille store Evenstad i en usikker verden. Full text in Research Archive
• Zimmermann, Barbara (2025). Elg i endring. Full text in Research Archive
• Zimmermann, Barbara (2025). GRENSEVILT - viltforvaltning på tvers. Full text in Research Archive
• Zimmermann, Barbara (2025). Store rovdyr i Nord-Gudbrandsdalen: Biologi, konflikter og forvaltning. Full text in Research Archive
• Zimmermann, Barbara (2025). Prosjektet "Beiting i Rovdyrskogen" oppsummert – de viktigste resultatene. Full text in Research Archive
• Zimmermann, Barbara; Hellbaum, Paige; Ausilio, Giorgia; Sand, Håkan; Wikenros, Camilla & Eriksen, Ane [Show all 11 contributors for this article] (2025). Counteracting management of a transboundary moose population in Scandinavia. Full text in Research Archive
• Zimmermann, Barbara (2025). Utmarksbeiting i rovdyrskogen. Full text in Research Archive
• Zimmermann, Barbara (2025). Utmarksbeiting i rovdyrskogen. Full text in Research Archive
• Zimmermann, Barbara; Forti, Alessandro; Lissillour, Pierre; Eriksen, Ane; Wabakken, Petter & Cerjak, Benoit [Show all 8 contributors for this article] (2025). Carnivore exclosures to protect sheep in the boreal forest affect the area use of moose. Full text in Research Archive
• Dyngeland, Cecilie; Zimmermann, Barbara & Mayer, Martin (2025). Storfe i rovdyrskogen: Utfordringer og muligheter. Full text in Research Archive
• Mayer, Martin; Dyngeland, Cecilie; Tofastrud, Morten; Wabakken, Petter & Zimmermann, Barbara (2025). Tap av storfe og sau i norsk utmark: hvorfor, hvor og når? Full text in Research Archive
• Zimmermann, Barbara; Forti, Alessandro; Lissillour, Pierre; Eriksen, Ane; Cerjak, Benoit & Campon, Carla [Show all 8 contributors for this article] (2025). Effekter av rovdyravvisende gjerder på storvilt. Full text in Research Archive
• Panzacchi, Manuela; Moorter, Bram Van; Niebuhr, Bernardo Brandão; Thorsen, Neri Horntvedt; Rolandsen, Christer Moe & Solberg, Erling Johan [Show all 10 contributors for this article] (2025). Kumulative effekter og habitatkonnektivitet: De første nasjonale kartene og deres bruk for å støtte bærekraftig forvaltning gjennom scenarioanalyser og sonering. Full text in Research Archive
• Versluijs, Erik; Zimmermann, Barbara; Niccolai, Laura Jeanne; Tofastrud, Morten; Hessle, Anna Karin & Scasta, John Derek [Show all 9 contributors for this article] (2025). How Advances in Livestock Technology Can Aid Wildlife Coexistence: a Case Study on Free-Ranging Cattle in the Boreal Forest. Full text in Research Archive Show summary

  In the last decade, technological advances have soared, both within wildlife research and livestock production. Within livestock farming, there has been a significant increase in technological applications to measure and monitor animal health, with the main focus on increasing production and maintaining high animal welfare standards. These precision livestock farming (PLF) techniques can be utilized beyond highly modernized barns. One such example is the use of accelerometers, i.e. activity sensors from which behaviours or certain activity states can be derived. For example, in an earlier study, we have successfully used accelerometers in free-ranging cattle in large, extensive forest areas in Norway to study cattle behaviours at high spatio-temporal resolution. However, interactions between livestock and wildlife bring multifaceted societal challenges of high complexity, such as livestock depredation. For example, repeated livestock depredation negatively affects animal welfare, is economically and emotionally straining to farmers, and leads to lower acceptance level for wildlife, such as carnivores, at a societal level. Studying wildlife species requires considerable effort, often using invasive techniques, and generally results in lower sample sizes due to lower population densities. Increasing focus on studying livestock within wildlife conflicts offers the opportunity to understand how different species, breeds, and age classes are affected by interactions with wildlife. At the same time, this provides valuable information about wildlife species. We have successfully used accelerometers on free-ranging cattle in large, extensive forest areas in Norway to study cattle behaviours at high spatio-temporal resolution. In our study, all adult cattle in the herds were equipped with virtual fence collars, recording both GPS and acceleration data. By using a behavioural classification model, we predicted behaviours and examined how different environmental, climatic, and topographic drivers relate to these behaviours. Ultimately, this can be used to study changes of behaviours, and potentially increased stress and vigilance behaviour in response to carnivore presence or other perceived risks. Furthermore, high livestock abundance and lower effort required for tagging livestock can increase the understanding of differences between breeds, ages, farming systems, etc., and therefore aid in coexistence with wildlife and reducing wildlife conflicts.

• Versluijs, Erik; Zimmermann, Barbara; Hessle, Anna Karin & Tofastrud, Morten (2025). Living with wolves: A new framework to enhance coexistence with wolves by studying livestock behaviours. Full text in Research Archive Show summary

  Coexisting with wolves brings multifaceted societal challenges of high complexity, such as livestock losses due to depredation. Countries often support farmers with economic compensation of livestock losses. However, farmers affected by repeated depredation events might not only suffer economically, but also emotionally. The resulting increase in stress and worry can impact their daily life, which in turn can negatively effect animal welfare. Furthermore, repeated livestock depredation will negatively effect animal welfare, as individuals which are not attacked who might have higher stress levels. In the meantime, many studies related to wolf management and coexistence are focussing directly on wolf behaviour and preventive measures to alleviate conflicts. These efforts are of great importance. However, in addition to this, we propose a new framework for studying livestock behaviours using precision livestock farming (PLF) techniques to enhance coexistence with wolves. Within livestock farming there is a strong increase in technological applications to measure and monitor animal health, with the main focus on increasing production and maintaining high animal welfare standards. These PLF techniques can reach further than only be used in highly modernized barns. One such example is the use of accelerometers, i.e. activity sensors from which behaviours or certain activity states can be derived. We have in an earlier study successfully used accelerometers in free-ranging cattle in large extensive forest areas in Norway to study cattle behaviours at high spatio-temporal resolution. Based on these experiences, we propose creating a broader network of cattle behaviour studies that consider different herd compositions, pasture types, breeds, ages, and management practices. Therefore, creating a wide knowledge base about breed specific livestock behaviours and responses towards wolf presence gives the opportunity to 1) inform farmers and suggest adjustments of management practices in specific situations, such as changing to breeds or herd compositions with a lower probability of depredation. And 2) developing tools to better monitor livestock behaviour, for example to detect sudden changes in behaviour suggesting towards wolf presence. In this presentation we will outline this interdisciplinary framework, discuss the technology and methods, and use a practical case to show how this might be implemented.

• Lorand, Charlotte Muguette Renée; Eriksen, Ane; Devineau, Olivier; Chimienti, Marianna; Wikenros, Camilla & Sand, Håkan [Show all 8 contributors for this article] (2025). Close encounters with wolves: who's tiptoeing around whom? Full text in Research Archive
• Zimmermann, Barbara (2024). Ulvens biologi og nyeste resultater fra ulveforskning. Full text in Research Archive
• Zimmermann, Barbara (2024). Viltforskning på Evenstad. Full text in Research Archive
• Zimmermann, Barbara (2024). Cross-border wolf recovery, management and research in Scandinavia. Full text in Research Archive
• Zimmermann, Barbara; Sand, Håkan; Ausilio, Giorgia; Wikenros, Camilla; Eriksen, Ane & Wabakken, Petter (2024). Ikke lett å være elg! Full text in Research Archive
• Zimmermann, Barbara (2024). Ulvens biologi som grunnlag for konflikter. Full text in Research Archive
• Eriksen, Ane; Lorand, Charlotte Muguette Renée; Versluijs, Erik; Chimienti, Marianna; Devineau, Olivier & Bouet, Lea [Show all 10 contributors for this article] (2024). Studying wolf flight responses during experimental approaches using fine-scale GPS and accelerometer data. Full text in Research Archive
• Eriksen, Ane; Zimmermann, Barbara; Kirchner, Theresa Margret; Evans, Alina & Mathisen, Karen Marie (2024). Elg i endring. Full text in Research Archive
• Eriksen, Ane; Lorand, Charlotte Muguette Renée; Versluijs, Erik; Chimienti, Marianna; Devineau, Olivier & Bouet, Lea [Show all 10 contributors for this article] (2024). Eksperimentelle nærmøter med ulv – oppdatering og framtidige planer. Full text in Research Archive
• Lorand, Charlotte Muguette Renée; Eriksen, Ane; Devineau, Olivier; Bouet, Lea; Chimienti, Marianna & Wikenros, Camilla [Show all 9 contributors for this article] (2024). The secret lives of wolves: Predicting behaviors in wild wolves approached by humans, using collar-mounted accelerometers and a model calibrated in captivity. Full text in Research Archive
• Zimmermann, Barbara (2024). Beskatning av elg fra rovdyr og mennesker i områder med bjørn og ulv. Full text in Research Archive
• Zimmermann, Barbara (2024). Grazing in carnivore forests for biodiversity and sustainable production of food and timber. Full text in Research Archive
• Zimmermann, Barbara (2024). Elgens dødelighet. Full text in Research Archive
• Zimmermann, Barbara (2024). Back in town: Wolf recovery in the multi-national landscape of Europe. Full text in Research Archive
• Zimmermann, Barbara (2024). Back in town: Wolf recovery in the multi-national landscape of Europe. Full text in Research Archive
• Zimmermann, Barbara (2024). Back in town: Wolf recovery in the multi-national landscape of Europe. Full text in Research Archive
• Zimmermann, Barbara (2024). Back in town: Wolf recovery in the multi-national landscape of Europe. Full text in Research Archive
• Zimmermann, Barbara; Hartog, Emma den; Dyngeland, Cecilie; Tofastrud, Morten; Wabakken, Petter & Kleiven, Jo (2024). Dødelighet og forebyggende tiltak mot rovviltskader hos storfe i Europa og Nord-Amerika 1999 - 2020. Full text in Research Archive
• Spedener, Mélanie; Hérault, Rémi; Hauer, Josh; Mathisen, Karen Marie; Tofastrud, Morten & Austrheim, Gunnar [Show all 7 contributors for this article] (2024). Positive and negative effects of cattle on small spruces in boreal production forest. Full text in Research Archive
• Kommisrud, Elisabeth & Zimmermann, Barbara (2024). From research idea to a finalized project. Full text in Research Archive
• Zimmermann, Barbara (2024). Rovdyr i grenseland. Full text in Research Archive
• Zimmermann, Barbara; Dyngeland, Cecilie; Mayer, Martin & Tofastrud, Morten (2024). Cattle farmers’ experiences with carnivores and dogs. Full text in Research Archive
• Zimmermann, Barbara (2024). GRENSEVILT – viltforvaltning på tvers. Full text in Research Archive
• Zimmermann, Barbara (2024). Wolf ecology and management in Norway. Full text in Research Archive
• Zimmermann, Barbara; Sand, Håkan; Ausilio, Giorgia; Wikenros, Camilla; Eriksen, Ane & Wabakken, Petter (2024). Ikke lett å være elg. Full text in Research Archive
• Zimmermann, Barbara (2024). Ett år med elgku E2102. Full text in Research Archive
• Zimmermann, Barbara; Niccolai, Laura Jeanne; Versluijs, Erik; Spedener, Mélanie; Evans, Alina & Tofastrud, Morten [Show all 7 contributors for this article] (2024). How can electronical tags for livestock facilitate livestock production in areas with large carnivores? Full text in Research Archive
• Zimmermann, Barbara (2024). Litt av hvert om ulven. Full text in Research Archive
• Zimmermann, Barbara (2024). Beiting i rovdyrskogen og nytt fra ulveforskningen. Full text in Research Archive
• Storch, Ilse; Tallian, Aimee Grace; Boitani, Luigi; Bernardi, Cecilia Di; Kusak, Josip & Şekercioğlu, Çağan [Show all 9 contributors for this article] (2024). Wolves across borders. Wildlife Biology. ISSN 0909-6396. 2024(6). doi: 10.1002/wlb3.01417. Full text in Research Archive
• Niccolai, Laura Jeanne; Wulff, Saskia H.; Versluijs, Erik; Spedener, Mélanie; Zimmermann, Barbara & Hessle, Anna [Show all 9 contributors for this article] (2024). Forest Cows Secrets: Cracking the Code With Movement Sensors. Frontiers for Young Minds. doi: 10.3389/frym.2024.1305706. Full text in Research Archive Show summary

  Have you ever wondered how we can watch animals in the wild without actually being near them? In Norway, cows roam freely in the deep forest during summer. While the cows enjoy the freedom, it can be tricky to keep them safe from carnivores like wolves and bears, as no shepherds or dogs protect the herds. Keeping an eye on the cows is important! Farmers and researchers use GPS to track animals, just as we do for phones or cars. However, GPS does not tell us much about what the animals are doing. That is where movement sensors come in. These sensors store information about the tiniest body movements and reveal what the animal is doing at any time. Is the cow’s head up or down? Is it walking or running? Based on the data, we could distinguish 20 different behaviors! Now we can spy on cows, see what they are up to in the forest, and help farmers better care for them.

• Versluijs, Erik; Zimmermann, Barbara; Tofastrud, Morten; Hessle, Anna; Scasta, John Derek & Serrouya, Robert [Show all 7 contributors for this article] (2024). Behavioural dependent habitat selection of free-ranging cattle in the boreal forest. Full text in Research Archive Show summary

  During the 20th century, when large carnivores were subject to eradication in Norway, farmers started to release their livestock unsupervised in the forests during summer. Since 1970, Norway has committed to conserve large carnivores, which led to high losses in domestic sheep herds. Therefore, farmers are encouraged to use cattle instead, as cattle have a hundred times lower depredation rate. However, farmers report increased stress in cattle, and income loss. We GPS collared all adult cattle (N = 125) in four herds and two areas in the Innlandet County of SE-Norway. The collars included accelerometer sensors and were equipped with virtual fencing technology to retain the animals inside areas up to 25 km2. We registered GPS positions of cattle using 15-minute intervals and predicted common behaviours such as grazing, walking, vigilance, and resting, using accelerometry signatures. We used multi-state step-selection analyses to study effects of environmental variables (habitat, weather) on cattle behaviours,body condition, and seasonal variability. Preliminary results indicated that cattle used mostly forest roads for walking, as previously shown. Our results provide novel insights about cattle’s space-use in boreal forests, and the potential to study the compatibility of free-ranging cattle with carnivores and effects on animal welfare.

• Mayer, Martin; Dyngeland, Cecilie & Zimmermann, Barbara (2024). Livestock losses due to large carnivores and other causes. Comparing Rovbase and OGA data with a farmer survey. Full text in Research Archive
• Zimmermann, Barbara (2024). Utmarksbeiting i rovdyrskogen for økt artsmangfold og bærekraftig produksjon av mat og tømmer. Full text in Research Archive
• Zimmermann, Barbara (2024). Elgens dødelighet i et risikofylt landskap. Full text in Research Archive
• Zimmermann, Barbara (2024). Ett år med elgku E2102. Full text in Research Archive
• Zimmermann, Barbara (2024). Dumb ways to die: Elgens dødelighet i et risikofylt landskap. Full text in Research Archive
• Adamski, Genevieve; Versluijs, Erik; Zimmermann, Barbara & Tofastrud, Morten (2023). Outstanding in their field: factors influencing cattle habitat selection in Norwegian summer ranging areas. Full text in Research Archive
• Talleraas, Simen Moflag; Midjaas, Tov; Zimmermann, Barbara; Mathisen, Karen Marie; Kalén, Christer & Pedersen, Simen (2023). Elg i et skiftende klima - hvordan påvirker snødekke elgens beitevalg om vinteren. Hjorteviltet. ISSN 1502-3729. p. 84–86. Full text in Research Archive
• Tallian, Aimee Grace; Ordiz, Andrés; Metz, Matthew; Zimmermann, Barbara; Wikenros, Camilla & Smith, Douglas [Show all 11 contributors for this article] (2023). Of wolves and bears: Seasonal drivers of interference and exploitation competition between apex predators. Full text in Research Archive
• Spedener, Mélanie & Zimmermann, Barbara (2023). Grazing in carnivore forests for sustainable production of food, timber and biodiversity. Full text in Research Archive
• Spedener, Mélanie; Mathisen, Karen Marie; Henriksen, Marie Vestergaard; Nielsen, Anders; Austrheim, Gunnar & Zimmermann, Barbara (2023). Effects of domestic animal grazing on biodiversity in forests. Full text in Research Archive
• Zimmermann, Barbara (2023). Cross-border wolf recovery, management and research in Scandinavia. Full text in Research Archive
• Tallian, Aimee Grace; Zimmermann, Barbara; Sand, Håkan; Wabakken, Petter & Wikenros, Camilla (2023). Wolves Across Borders: Conference on Wolf Ecology and Management. Full text in Research Archive
• Kalén, Christer; Zimmermann, Barbara; Pedersen, Simen; Sjølie, Hanne Kathrine; Dyngeland, Cecilie & Franklin, Oskar (2023). Forest and wildlife under pressure - sustainable analysis for sustainable solutions. Full text in Research Archive
• Zimmermann, Barbara; Mathisen, Karen Marie; Wikenros, Camilla & Sand, Håkan (2023). Elgforvaltning i grenseland. Full text in Research Archive
• Zimmermann, Barbara (2023). Management of cross-bordering wildlife. Full text in Research Archive
• Zimmermann, Barbara (2023). GRENSEVILT – viltforvaltning på tvers. Full text in Research Archive
• Bernardi, Cecilia Di; Åkesson, Mikael; Sand, Håkan; Aronsson, Malin; Ciucci, Paolo & Boitani, Luigi [Show all 12 contributors for this article] (2023). Linking wolf feeding behavior to individual predator traits. Full text in Research Archive
• Niccolai, Laura Jeanne; Ausilio, Giorgia; Wikenros, Camilla; Sand, Håkan; Wabakken, Petter & Devineau, Olivier [Show all 7 contributors for this article] (2023). Dancing with Wolves: Behavioral responses of moose to close proximity of wolves in Scandinavia. Full text in Research Archive
• Hartog, Emma den; Dyngeland, Cecilie; Tofastrud, Morten; Kleiven, Jo; Scasta, John Derek & Serrouya, Robert [Show all 7 contributors for this article] (2023). Carnivore depredation on cattle and measures to prevent this: Comparing experiences across Europe and North America. Full text in Research Archive
• Ausilio, Giorgia; Wikenros, Camilla; Sand, Håkan; Devineau, Olivier; Eriksen, Ane & Wabakken, Petter [Show all 10 contributors for this article] (2023). Environmental and anthropogenic features mediate human hunting risk and wolf predation risk for moose. Full text in Research Archive
• Zimmermann, Barbara (2023). Applications in the borderland. Full text in Research Archive
• Versluijs, Erik; Zimmermann, Barbara; Tofastrud, Morten; Hessle, Anna; Evans, Alina & Adamski, Genevieve [Show all 9 contributors for this article] (2023). Towards a shared forest: Studying the compatibility of free-ranging cattle and carnivores in boreal forests. Full text in Research Archive
• Zimmermann, Barbara; Ausilio, Giorgia; Sand, Håkan; Wikenros, Camilla; Eriksen, Ane & Wabakken, Petter [Show all 7 contributors for this article] (2023). Moose at risk: Scandinavian biotagging research on impacts of predation, harvest and climate change. Full text in Research Archive
• Zimmermann, Barbara; Wikenros, Camilla; Mathisen, Karen Marie; Ausilio, Giorgia; Sand, Håkan & Hellbaum, Paige [Show all 12 contributors for this article] (2023). Elgforvaltning i grenseland. Full text in Research Archive
• Zimmermann, Barbara (2023). GRENSEVILT – viltforvaltning på tvers. Full text in Research Archive
• Mayer, Martin; Furuhovde, Erlend; Nordli, Kristoffer; Ausilio, Giorgia; Eriksen, Ane & Evans, Alina [Show all 8 contributors for this article] (2023). Efficiency and disturbance effects of drone versus ground approaches to monitor moose. Full text in Research Archive
• Ausilio, Giorgia; Sand, Håkan; Wikenros, Camilla; Zimmermann, Barbara; Wabakken, Petter & Mathisen, Karen Marie (2023). Jakt- och vargpredationsrisk påverkar älgens habitatval. Fakta Skog. ISSN 1400-7789. 2023-5. Full text in Research Archive
• Sand, Håkan; Liberg, Olof; Wikenros, Camilla; Åkesson, Mikael; Zimmermann, Barbara & Wabakken, Petter (2023). High turnover rates of territories in the Scandinavian wolf population: causes and consequences. Full text in Research Archive
• Zimmermann, Barbara; Sand, Håkan; Dyngeland, Cecilie & Lund, Signe Sannem (2023). Elgjakt i en skog av vindturbiner. Hjorteviltet. ISSN 1502-3729. Full text in Research Archive
• Ausilio, Giorgia; Sand, Håkan; Wikenros, Camilla; Aronsson, Malin; Milleret, Cyril Pierre & Nordli, Kristoffer [Show all 11 contributors for this article] (2023). Effects of large carnivores, hunter harvest and climate on the survival of calves in a partially migratory moose population. Full text in Research Archive
• Zimmermann, Barbara; Wabakken, Petter & Eriksen, Ane (2023). Carnivore exclosures to protect sheep: Experiences and effects on wildlife. Full text in Research Archive
• Zimmermann, Barbara (2023). Ikke lett å være elg. Full text in Research Archive
• Spedener, Mélanie; Nielsen, Anders; Henriksen, Marie Vestergaard; Austrheim, Gunnar; Mathisen, Karen Marie & Zimmermann, Barbara (2023). Cattle, wild ungulates and the plant- pollinator community in young spruce plantations. Full text in Research Archive
• Zimmermann, Barbara; Lund, Signe Sannem; Dyngeland, Cecilie; Sand, Håkan & Wikenros, Camilla (2023). Elgjakt ved vindkraftverk. Full text in Research Archive
• Zimmermann, Barbara; Ausilio, Giorgia; Sand, Håkan; Wikenros, Camilla; Eriksen, Ane & Wabakken, Petter [Show all 7 contributors for this article] (2023). Moose at risk: Scandinavian biotagging research on impacts of predation, harvest and climate change. Full text in Research Archive
• Zimmermann, Barbara (2023). Norsk og svensk beitetakstmetodikk. Full text in Research Archive
• Zimmermann, Barbara & Ausilio, Giorgia (2023). 37th IUGB Congress 2025: How wild is wildlife? Full text in Research Archive
• Falkevik, Maria; Esselin, Anders; Zimmermann, Barbara & Wikenros, Camilla (2023). Grensevilt: Proposal of a cooperation platform for cross-border wildlife management in the Nordic countries. Full text in Research Archive
• Wikenros, Camilla & Zimmermann, Barbara (2023). The Swedish-Norwegian border causes source-sink dynamics for wildlife but results in a win-win for research. Full text in Research Archive
• Zimmermann, Barbara (2023). GRENSEVILT – viltforvaltning på tvers. Full text in Research Archive
• Zimmermann, Barbara (2023). Norges store rovdyr. Full text in Research Archive
• Zimmermann, Barbara (2023). Norsk og svensk beitetakstmetodikk gir motsatt utslag i elgforvaltningen. Full text in Research Archive
• Zimmermann, Barbara (2023). Ulvens biologi og nytt fra ulveforskningen. Full text in Research Archive
• Zimmermann, Barbara (2023). Tiltak for å minske beiteskader. Full text in Research Archive
• Zimmermann, Barbara (2023). GRENSEVILT – viltforvaltning på tvers. Full text in Research Archive
• Zimmermann, Barbara (2023). Norsk og svensk beitetakstmetodikk gir motsatt utslag i elgforvaltningen. Full text in Research Archive
• Zimmermann, Barbara (2023). Elg i endring: Predasjon, jakt og klimaendringer. Full text in Research Archive
• Zimmermann, Barbara; Mathisen, Karen Marie; Hellbaum, Paige; Ausilio, Giorgia; Sand, Håkan & Wikenros, Camilla [Show all 12 contributors for this article] (2023). Contrasting management goals for a transboundary moose population in Scandinavia. Full text in Research Archive
• Zimmermann, Barbara (2023). Beitetakst på norsk og svensk vis: Motsatte konklusjoner for felles elgbestand. Full text in Research Archive
• Zimmermann, Barbara (2023). Elg i endring: Predasjon, jakt og klimaendringer. Full text in Research Archive
• Zimmermann, Barbara (2023). Grazing in carnivore forests for sustainable production of food, timber and biodiversity. Full text in Research Archive
• Ausilio, Giorgia; Zimmermann, Barbara; Mathisen, Karen Marie; Eriksen, Ane; Wabakken, Petter & Wikenros, Camilla [Show all 7 contributors for this article] (2023). Effekter av varg och andra faktorer på älgförekomst och betesskador. Full text in Research Archive
• Milleret, Cyril Pierre; Dupont, Pierre; Dey, Soumen; Broseth, Henrik; Kindberg, Jonas & Turek, Daniel [Show all 11 contributors for this article] (2023). Mapping population level mortality risk of wolves in Scandinavia. Full text in Research Archive
• Milleret, Cyril Pierre; Dupont, Pierre; Dey, Soumen; Broseth, Henrik; Kindberg, Jonas & Turek, Daniel [Show all 11 contributors for this article] (2023). Mapping population level mortality risks of an emblematic and controversial large carnivore. Full text in Research Archive
• Tallian, Aimee Grace; Zimmermann, Barbara; Kindberg, Jonas; Ordiz, Andres; Metz, Matthew C. & Wikenros, Camilla [Show all 11 contributors for this article] (2022). Sympatric predators: The effect of brown bears on wolf hunting behavior. Full text in Research Archive
• Zimmermann, Barbara (2022). Ujevn romlig fordeling av ulvens predasjon på elg. Full text in Research Archive
• Zimmermann, Barbara; Sand, Håkan; Wikenros, Camilla; Eriksen, Ane & Wabakken, Petter (2022). Uneven spatial and temporal distribution of predation inside wolf territories challenges moose management. Full text in Research Archive
• Zimmermann, Barbara (2022). Wolf control in Scandinavia. Full text in Research Archive
• Zimmermann, Barbara (2022). Familieidyll med spisse tenner: Ulvens biologi som grunnlag for interessekonflikter. Full text in Research Archive
• Zimmermann, Barbara (2022). Resultater fra GRENSEVILT 1 og 2. Full text in Research Archive
• Zimmermann, Barbara & Wahl, Andreas (2022). Folkeopplysningen: Ulv. [TV]. NRK. Full text in Research Archive
• Zimmermann, Barbara (2022). LARGE research group management. Full text in Research Archive
• Zimmermann, Barbara (2022). Wolf population regulation in Scandinavia. Full text in Research Archive
• Zimmermann, Barbara (2022). Wolf population management in Scandinavia. Full text in Research Archive
• Zimmermann, Barbara; Wikenros, Camilla; Aronsson, Malin; Eriksen, Ane; Mathisen, Karen Marie & Persson, Jens [Show all 8 contributors for this article] (2022). Viltforvaltning på tvers av grensen: Ny lærdom fra GRENSEVILT prosjektet. Jakt og Fiske i Hedmark. ISSN 1891-1471. Full text in Research Archive
• Sand, Håkan; Zimmermann, Barbara; Wikenros, Camilla & Wabakken, Petter (2022). Ulvens påvirkning på elg i en fortettet ulvebestand. Våre Rovdyr. ISSN 0801-4728. 3. Full text in Research Archive
• Sand, Håkan; Zimmermann, Barbara; Berg, Erik; Bramorska, Beata; Eriksen, Ane & Wikenros, Camilla [Show all 10 contributors for this article] (2022). Trekkmønster hos grenseelg – konsekvenser for forvaltningen. Hjorteviltet. ISSN 1502-3729. Full text in Research Archive
• Zimmermann, Barbara (2022). GRENSEVILT – viltforvaltning på tvers. Full text in Research Archive
• Zimmermann, Barbara; Mathisen, Karen Marie; Ausilio, Giorgia; Sand, Håkan; Wikenros, Camilla & Eriksen, Ane [Show all 13 contributors for this article] (2022). Motsatte norske og svenske mål for felles elgbestand. Hjorteviltet. ISSN 1502-3729. Full text in Research Archive
• Zimmermann, Barbara (2022). Large carnivores of Scandinavia. Full text in Research Archive
• Zimmermann, Barbara (2022). Beitetakst på norsk og svensk vis: Motsatte konklusjoner for felles elgbestand. Full text in Research Archive
• Zimmermann, Barbara (2022). Status for prosjektene GRENSEVILT og CarniForeGraze. Full text in Research Archive
• Zimmermann, Barbara (2022). Beitetakst på norsk og svensk vis. Full text in Research Archive
• Zimmermann, Barbara (2022). Familieidyll med spisse tenner: Ulvebiologi som grunnlag for interessekonflikter. Full text in Research Archive
• Tallian, Aimee Grace; Zimmermann, Barbara; Sand, Håkan; Wabakken, Petter & Wikenros, Camilla (2022). Wolves Across Borders: Conference on Wolf Ecology and Management. Full text in Research Archive
• Zimmermann, Barbara (2022). Resultater fra GRENSEVILT. Full text in Research Archive
• Zimmermann, Barbara (2022). Big teeth, big wounds: Large carnivores in Scandinavia. Full text in Research Archive
• Zimmermann, Barbara (2022). Elgens påvirkning på skog i grenseregionen. Full text in Research Archive
• Zimmermann, Barbara (2022). Ulvens biologi og ny forskning. Full text in Research Archive
• Zimmermann, Barbara (2022). Ny forskning om ulvens påvirkning på elg og elgjakt. Full text in Research Archive
• Zimmermann, Barbara (2022). Elgtrekk på tvers av grensen: påvirkning på ulv, skog og jakt. Full text in Research Archive
• Zimmermann, Barbara (2022). Management of cross-bordering wildlife. Full text in Research Archive
• Zimmermann, Barbara (2022). Elgvandringer i grenseland med følger for skogbruk og jakt. Full text in Research Archive
• Talleraas, Simen Moflag; Kalén, Crister; Zimmermann, Barbara; Franklin, Oskar; Evans, Alina & Eriksen, Ane [Show all 7 contributors for this article] (2022). How does bilberry affect the availability and use of bilberry forage for moose? Full text in Research Archive
• Tallian, Aimee Grace; Ordiz, Andrés; Metz, Matthew; Zimmermann, Barbara; Wikenros, Camilla & Smith, Douglas [Show all 11 contributors for this article] (2021). Sympatric predators: The effect of brown bears on wolf hunting behavior. Full text in Research Archive
• Tallian, Aimee Grace; Ordiz, Andrés; Metz, Matthew; Zimmermann, Barbara; Wikenros, Camilla & Smith, Douglas [Show all 11 contributors for this article] (2021). Wolves and bears: The nature of competitive interactions. Full text in Research Archive
• Zimmermann, Barbara (2021). Forebyggende tiltak mot rovviltskader. Full text in Research Archive
• Zimmermann, Barbara (2021). Predasjon generelt og resultater fra forskning på ulv og elg. Full text in Research Archive
• Løberg, Anne Kari; Zimmermann, Barbara & Gangås, Kristin Evensen (2021). Tillitt til rovviltforskning. [Radio]. NRK P2 Nyhetsmorgen. Full text in Research Archive
• Zimmermann, Barbara; Gangås, Kristin Evensen & Løberg, Anne Kari (2021). Ulveforskere blir motarbeida og ikke trodd på. [Internet]. www.nrk.no. Full text in Research Archive
• Ringlund, Ole Martin; Zimmermann, Barbara; Gangås, Kristin Evensen & Strand, Mona (2021). Forskningsdagene: Rovdyrkonflikten. [Internet]. https://www.youtube.com/watch?v=WoIHCGZ2R28. Full text in Research Archive
• Zimmermann, Barbara & Flagstad, Øystein (2021). Tetthet og innavl påvirker når ulv i Skandinavia får sine første valpekull. [Journal]. Nationen. Full text in Research Archive
• Zimmermann, Barbara & Hystad, Joar (2021). Lav tillit til forskning på ulv og andre rovdyr: — Folk har sterke meninger. [Internet]. Khrono.no. Full text in Research Archive
• Zimmermann, Barbara & Kjørstad, Elise (2021). Du har kanskje hørt om alfahannen og alfahunnen i en ulveflokk? Begrepet er basert på en misforståelse, ifølge eksperter. [Internet]. forskning.no. Full text in Research Archive
• Zimmermann, Barbara & Nordli, Kristoffer (2021). Wildnis Europa Vielfrass. [TV]. ARTE TV. Full text in Research Archive
• Zimmermann, Barbara & Kjørstad, Elise (2021). Wolf packs don’t actually have alpha males and alpha females, the idea is based on a misunderstanding. [Internet]. sciencenorway.no. Full text in Research Archive
• Zimmermann, Barbara; Flagstad, Øystein & Stokmo, Jan Arne (2021). Tetthet og innavl påvirker når ulv i Skandinavia får sine første valpekull. [Internet]. forskning.no. Full text in Research Archive
• Zimmermann, Barbara (2021). Beitingas påvirkning på viltet. Full text in Research Archive
• Zimmermann, Barbara & Vogler, Thomas (2021). Drone field demonstrations in Norway. Full text in Research Archive
• Zimmermann, Barbara (2021). Anvendt økologisk forskning på Evenstad. Full text in Research Archive
• Zimmermann, Barbara (2021). Ulvens betydning for elgbestanden og elgjakta. Full text in Research Archive
• Zimmermann, Barbara (2021). Ulvens betydning for elgbestanden og elgjakta. Full text in Research Archive
• Zimmermann, Barbara (2021). Prosjektet Grensevilt og studier på elgtrekk. Full text in Research Archive
• Zimmermann, Barbara (2021). Wolf effects on moose and moose harvest. Full text in Research Archive
• Zimmermann, Barbara (2021). Beitingas påvirkning på viltet. Full text in Research Archive
• Zimmermann, Barbara (2021). Ulvepredasjon i Skandinavia. Full text in Research Archive
• Zimmermann, Barbara (2021). Drone pilotprosjekt med GRENSEVILT-elg. Full text in Research Archive
• Zimmermann, Barbara (2021). Elgvandringer i grenseland. Full text in Research Archive
• Zimmermann, Barbara (2021). Utmarksbeiting i rovdyrskogen for bærekraftig produksjon av mat og tømmer og økt artsmangfold. Full text in Research Archive
• Zimmermann, Barbara (2021). Resultater fra GRENSEVILT: Predasjon på elg i en fortettet ulvebestand. Full text in Research Archive
• Zimmermann, Barbara (2021). Rovdyrene i Norge. Full text in Research Archive
• Wikenros, Camilla; Zimmermann, Barbara; Eriksen, Ane; Falkevik, Maria; Lund, Pia K. & Løken, Øivind [Show all 10 contributors for this article] (2021). GRENSEVILT 2 – forvaltningssamarbeit og effekter av vindkraft. Hjorteviltet. ISSN 1502-3729. Full text in Research Archive
• Zimmermann, Barbara (2021). Rovdyrenes biologi. Full text in Research Archive
• Zimmermann, Barbara (2021). Rovvilt: økologi, konflikter og forvaltning. Full text in Research Archive
• Wikenros, Camilla; Zimmermann, Barbara; Eriksen, Ane; Persson, Jens; Sand, Håkan & Wabakken, Petter (2021). GRENSEVILT 2 – forvaltningssamarbeid og effekter av vindkraft. Hjorteviltet. ISSN 1502-3729. Full text in Research Archive
• Zimmermann, Barbara (2021). Management of cross-bordering wildlife. Full text in Research Archive
• Zimmermann, Barbara (2021). Forvaltning av trekkelg på tvers av landegrenser – nytt fra GRENSEVILT. Full text in Research Archive
• Zimmermann, Barbara (2021). GRENSEVILT - viltforvaltning på tvers av grensen. Full text in Research Archive
• Wikenros, Camilla; Månsson, Johan; Maartmann, Erling; Eriksen, Ane; Wabakken, Petter & Zimmermann, Barbara (2021). Publisher Correction: Impact of a recolonizing, cross-border carnivore population on ungulate harvest in Scandinavia (Scientific Reports, (2020), 10, 1, (21670), 10.1038/s41598-020-78585-8). Scientific Reports. 11(1). doi: 10.1038/s41598-021-89263-8. Full text in Research Archive
• Tallian, Aimee Grace; Zimmermann, Barbara; Sand, Håkan; Wabakken, Petter & Wikenros, Camilla (2021). Wolves Across Borders: Conference on Wolf Ecology and Management. Full text in Research Archive
• Tofastrud, Morten & Zimmermann, Barbara (2021). Bruk av utmarksbeite til storfe. Buskap : medlemsblad for Geno. ISSN 0807-5069. p. 28–30. Full text in Research Archive
• Spedener, Mélanie; Nielsen, Anders; Henriksen, Marie Vestergaard; Austrheim, Gunnar; Mathisen, Karen Marie & Zimmermann, Barbara (2021). Does cattle grazing in young spruce plantations affect flowering plant diversity? Full text in Research Archive
• Wikenros, Camilla; Zimmermann, Barbara; Sand, Håkan; Månsson, Johan; Maartmann, Erling & Eriksen, Ane [Show all 7 contributors for this article] (2021). Så påverkar vargen älgobs och avskjutning. Svensk Jakt. ISSN 0039-6583. p. 30–32. Full text in Research Archive
• Zimmermann, Barbara (2020). Utmarksbeiting i rovdyrskogen for bærekraftig produksjon av mat og tømmer og økt artsmangfold. Full text in Research Archive
• Zimmermann, Barbara (2020). Nytt forskningsprosjekt om utmarksbeiting i rovdyrskogen. Full text in Research Archive
• Zimmermann, Barbara & Nyberget, Mari (2020). Forskningsprosjektet GRENSEVILT 2. [Radio]. Lokalradioene i Innlandet. Full text in Research Archive
• Zimmermann, Barbara (2020). Nytt fra ulveforskningen. Full text in Research Archive
• Zimmermann, Barbara (2020). GRENSEVILT 1: Wolf predation in change. Full text in Research Archive
• Zimmermann, Barbara & Falkevik, Maria (2020). Forskning och förvaltning: självständiga singlar eller det trogna, strävsamma paret? Full text in Research Archive
• Zimmermann, Barbara (2020). Ulvebiologi. Full text in Research Archive
• Zimmermann, Barbara (2020). Predasjon generelt og ulvepredasjon. Full text in Research Archive
• Zimmermann, Barbara (2020). Jakten på elg og kunnskap. Full text in Research Archive
• Zimmermann, Barbara & Nyberget, Mari (2020). Skal kartlegge hvordan vindkraftverket i Kjølberget pavirker-vilt. [Internet]. https://www.isolor.no/2020/nyheter. Full text in Research Archive
• Zimmermann, Barbara & Løberg, Anne Kari (2020). Norges mest ulvetette by. [Internet]. https://www.nrk.no/innlandet/norges-mest-ulvetette-by-1.1500. Full text in Research Archive
• Wikenros, Camilla; Zimmermann, Barbara; Sand, Håkan; Månsson, Johan; Maartmann, Erling & Eriksen, Ane [Show all 7 contributors for this article] (2020). Sett og felt elg i forhold til ulveforekomst i Norge og Sverige. Hjorteviltet. ISSN 1502-3729. Full text in Research Archive
• Zimmermann, Barbara (2020). Ulvens betydning for elg og elgjakt. Full text in Research Archive
• Zimmermann, Barbara (2020). Økologisk forskning på Evenstad. Full text in Research Archive
• Zimmermann, Barbara & Næsheim, Anne (2020). Stor elgmøkktelling skal gi bedre forvaltning av elgstammen. [TV]. https://www.nrk.no/innlandet/stor-elgmokktelling-skal-gi-bed. Full text in Research Archive
• Zimmermann, Barbara (2020). GRENSEVILT in a nutshell. Full text in Research Archive
• Zimmermann, Barbara (2020). Grensevilt og merking av dyr. Full text in Research Archive
• Zimmermann, Barbara; Evans, Alina; Wabakken, Petter; Mathisen, Karen Marie & Sjølie, Hanne Kathrine (2020). Utmarksbeiting i rovdyrskogen for bærekraftig produksjon av mat og tømmer og økt artsmangfold. Full text in Research Archive
• Tallian, Aimee Grace; Zimmermann, Barbara; Sand, Håkan; Wabakken, Petter & Wikenros, Camilla (2020). Wolves Across Borders: Conference on Wolf Ecology and Management. Full text in Research Archive
• Zimmermann, Barbara; Sand, Håkan; Wikenros, Camilla; Eriksen, Ane & Wabakken, Petter (2020). Å dele utbyttet med ulven: Jakt- og predasjonstrykk på elg i ulverevir i Skandinavia. Hjorteviltet. ISSN 1502-3729. Full text in Research Archive
• Zimmermann, Barbara; Sand, Håkan; Wikenros, Camilla; Eriksen, Ane & Wabakken, Petter (2020). Ujevn fordeling av ulvens uttak av elg i ulverevir påvirker jaktuttaket lokalt. Hjorteviltet. ISSN 1502-3729. Full text in Research Archive
• Zimmermann, Barbara (2020). Beskatning av elg i ulverevir. Full text in Research Archive
• Zimmermann, Barbara (2020). Resultater fra Grenseviltprosjektet. Full text in Research Archive
• Tallian, Aimee Grace; Ordiz, Andrés; Zimmermann, Barbara; Sand, Håkan; Wikenros, Camilla & Wabakken, Petter [Show all 8 contributors for this article] (2019). The relative effect of large carnivores and climate on ungulate productivity. Full text in Research Archive
• Eriksen, Ane; Zimmermann, Barbara; Wikenros, Camilla; Mathisen, Karen Marie; Wabakken, Petter & Sand, Håkan [Show all 8 contributors for this article] (2019). Grensevilt – reduksjon av grensebarrierer for skandinavisk viltforvaltning. Hjorteviltet. ISSN 1502-3729. Full text in Research Archive
• Zimmermann, Barbara & Wabakken, Petter (2019). GRENSEVILT: Svensk-norsk forskningsprosjekt om viltforvaltning tvers av grenser. Full text in Research Archive
• Devineau, Olivier; Zimmermann, Barbara; Cromsigt, Joris P.G.M.; Pfeffer, Sabine; Skarpe, Christina & Mathisen, Karen Marie (2019). Roads, forestry, and wolves interact to drive moose browsing behavior in Scandinavia. Full text in Research Archive
• Wikenros, Camilla & Zimmermann, Barbara (2019). Effekter av varg på avskjutning och älgobs. Full text in Research Archive
• Zimmermann, Barbara (2019). Ulvens påvirkning på elg og elgforvaltning. Full text in Research Archive
• Mathisen, Karen Marie; Sand, Håkan; Ausilio, Giorgia; Gicquel, M; Wikenros, Camilla & Månsson, Johan [Show all 9 contributors for this article] (2019). Trophic cascades in anthropogenic landscapes - Can recolonizing wolves reduce moose browsing and increase biodiversity? Full text in Research Archive
• Tallian, Aimee Grace; Ordiz, Andrés; Zimmermann, Barbara; Sand, Håkan; Wikenros, Camilla & Wabakken, Petter [Show all 8 contributors for this article] (2019). The relative effect of large carnivores and climate on ungulate productivity. Full text in Research Archive
• Eriksen, Ane; Versluijs, Erik; Fuchs, Boris; Zimmermann, Barbara; Wabakken, Petter & Bouet, Lea [Show all 16 contributors for this article] (2019). A standardized method for conducting and analysing experimental human approaches on wolves. Full text in Research Archive
• Loosen, Anne Elizabeth; Devineau, Olivier; Zimmermann, Barbara; Cromsigt, Joris P. G. M.; Skarpe, Christina & Pfeffer, Sabine [Show all 7 contributors for this article] (2019). Roads, forestry, and wolves interact to drive moose browsing behavior in Scandinavia. Full text in Research Archive
• Zimmermann, Barbara (2019). Research on large carnivores and moose, Grensevilt. Full text in Research Archive
• Zimmermann, Barbara (2019). Carnivore conflicts in Scandinavia. Full text in Research Archive
• Zimmermann, Barbara (2019). Jerven er tilbake i barskogen. Full text in Research Archive
• Zimmermann, Barbara (2019). Ulv gjennom et helt år. Full text in Research Archive
• Zimmermann, Barbara (2019). Ny forskning på forvaltning av elg i områder med ulv. Full text in Research Archive
• Zimmermann, Barbara (2019). Ulvens påvirkning på elg og elgforvaltning. Full text in Research Archive
• Zimmermann, Barbara (2019). Wolf predation research in GRENSEVILT. Full text in Research Archive
• Zimmermann, Barbara (2019). Ulvesona fra et forskningståsted og kort presentasjon av GRENSEVILT. Full text in Research Archive
• Zimmermann, Barbara (2019). Ulvens påvirkning på elg, jakt og forvaltning. Full text in Research Archive
• Zimmermann, Barbara (2019). Ulvens påvirkning på elg og elgforvaltning. Full text in Research Archive
• Zimmermann, Barbara (2019). Ulvens betydning for elg og elgjakt. Full text in Research Archive
• Zimmermann, Barbara (2019). Ulvens betydning for elg og elgjakt. Full text in Research Archive
• Zimmermann, Barbara (2019). Forskning på rovdyr og klauvvilt. Full text in Research Archive
• Zimmermann, Barbara (2019). Ulvens betydning for elg, Grensevilt. Full text in Research Archive
• Zimmermann, Barbara; Sand, Håkan; Wikenros, Camilla; Wabakken, Petter & Eriksen, Ane (2019). The asymmetric share of moose as prey for humans and wolves. Full text in Research Archive
• Zimmermann, Barbara (2019). Ulvens betydning for elg. Full text in Research Archive
• Zimmermann, Barbara; Breisjøberget, Jo Inge; Farstad, Espen & Skillingstad, Trond Gunnar (2019). Ulv!Ulv! [Internet]. https://www.statskog.no/jakt-og-fiske/podcast. Full text in Research Archive
• Zimmermann, Barbara & Wikenros, Camilla (2019). Kommunikasjonsarbeid i Grensevilt. Full text in Research Archive
• Zimmermann, Barbara (2019). Ulvens betydning for elg og elgjakt. Full text in Research Archive
• Zimmermann, Barbara & Falkevik, Maria (2019). Research and management: independent singles, happily married, sugaring or prostitution? Full text in Research Archive
• Zimmermann, Barbara (2019). Grensevilt - en presentasjon av prosjektet. Full text in Research Archive
• Zimmermann, Barbara; Breisjøberget, Jo Inge; Farstad, Espen & Skillingstad, Trond Gunnar (2019). Grenseløs forvaltning. [Internet]. https://www.statskog.no/jakt-og-fiske/podcast. Full text in Research Archive
• Zimmermann, Barbara (2019). Drones for wildlife management in forested areas. Full text in Research Archive
• Eriksen, Ane; Zimmermann, Barbara; Sunde, Peter; Gorini, Lucrezia; Wabakken, Petter & Evans, Alina [Show all 13 contributors for this article] (2018). Wolf responses to experimental approaches by humans along a gradient of anthropogenic impact: A pilot study. Full text in Research Archive
• Eriksen, Ane; Zimmermann, Barbara; Wabakken, Petter; Sanchez, David Carricondo; Sand, Håkan & Wikenros, Camilla (2018). Wolf movement relative to human structures during natal dispersal. Full text in Research Archive
• Zimmermann, Barbara (2018). Ulvers bevegelser i forhold til menneskers bosetting og aktivitet. Full text in Research Archive
• Zimmermann, Barbara & Wikenros, Camilla (2018). GRENSEVILT: Svensk-norskt forskningsprojekt om viltförvaltning över gränsen. Full text in Research Archive
• Zimmermann, Barbara (2018). Ulvens påvirkning på elg og elgforvaltning. Full text in Research Archive
• Zimmermann, Barbara; Wabakken, Petter; Eriksen, Ane; Mathisen, Karen Marie; Wikenros, Camilla & Sand, Håkan [Show all 8 contributors for this article] (2018). GRENSEVILT. Reduksjon av grensebarrierer for skandinavisk viltforvaltning. Interreg-prosjekt 2018-2020. Full text in Research Archive
• Zimmermann, Barbara; Wabakken, Petter; Eriksen, Ane; Sanchez, David Carricondo; Versluijs, Erik & Sand, Håkan [Show all 7 contributors for this article] (2018). Wolf tracks at the doorstep: A one-year cycle of wolf behavior close to houses in Scandinavia. Full text in Research Archive
• Zimmermann, Barbara (2018). Hva gjør ulven så politisk? Full text in Research Archive
• Zimmermann, Barbara (2018). Rovdyrforskning på Evenstad. Full text in Research Archive
• Zimmermann, Barbara (2018). Ulveatferd som byr på forvaltningsutfordringer. Full text in Research Archive
• Gorini, Lucrezia & Zimmermann, Barbara (2018). Managing large carnivores: the case of the wolf (Canis lupus) in Italy and Norway. Scienza Senza Confini. p. 27–31. Full text in Research Archive
• Zimmermann, Barbara (2018). Grensevilt, et nytt Interregprosjekt. Full text in Research Archive
• Zimmermann, Barbara (2018). Vilt i grenseland. Full text in Research Archive
• Zimmermann, Barbara (2018). Impact of wolf predation on moose. Full text in Research Archive
• Zimmermann, Barbara (2018). Ulv og nærhet til mennesker. Full text in Research Archive
• Zimmermann, Barbara (2018). Ulvens påvirkning på elg. Full text in Research Archive
• Zimmermann, Barbara & Wikenros, Camilla (2018). Grensevilt - et nytt Interregprosjekt. Full text in Research Archive
• Zimmermann, Barbara (2018). Storfe som alternativ til sau i rovdyrutsatte områder. Full text in Research Archive
• Zimmermann, Barbara & Lund, Frank Robert (2018). Brobyggeren på Evenstad. [Journal]. Hjorteviltet. Full text in Research Archive
• Zimmermann, Barbara (2018). Nytt fra ulveforskningen. Full text in Research Archive
• Zimmermann, Barbara (2018). GRENSEVILT - Reduksjon av grensebarrierer for skandinavisk viltforvaltning. Full text in Research Archive
• Zimmermann, Barbara (2018). Cats of the research group APEX. Full text in Research Archive
• Zimmermann, Barbara (2018). Grensevilt - en innføring i prosjektet. Full text in Research Archive
• Zimmermann, Barbara (2018). Wolf predation research. Full text in Research Archive
• Wabakken, Petter; Maartmann, Erling; Eriksen, Ane; Zimmermann, Barbara; Flagstad, Øystein & Liberg, Olof [Show all 8 contributors for this article] (2018). Betydningen av ulvens spredningsmønster for tap av bufe, tamrein og hund i Norge. Full text in Research Archive
• Sand, Håkan; Eklund, Ann; Wikenros, Camilla; Zimmermann, Barbara & Wabakken, Petter (2018). Vargens val: älg eller rådjur? Svensk Jakt. ISSN 0039-6583. 2(3), p. 34–37. Full text in Research Archive
• Nordli, Kristoffer; Zimmermann, Barbara; Wabakken, Petter & Eriksen, Ane (2018). On the way to independence - Ebbing cohesion in Scandinavian wolf family groups. Full text in Research Archive
• Loosen, Anne Elizabeth; Joris, Cromsigt; Zimmermann, Barbara; Devineau, Olivier & Mathisen, Karen Marie (2018). Do wolves influence moose browsing behavior in relation to roads in Scandinavia? Full text in Research Archive Show summary

  Wolves (Canis lupus) were functionally extinct in southern Fennoscandia by the mid-20th century, but are now recolonizing portions of their former range. Moose (Alces alces) are the primary prey of wolves. Moose, wolves and humans use low-traffic, secondary roads, and road-sides provide high-quality and quantity browse for moose. Roads alter predator-prey relationships by increasing predator movements, encounter rates with prey, and prey vigilance levels, thus contributing to a “landscape of fear”. Foraging theory predicts that moose will respond to riskier landscape by selecting habitats that reduce predation risk, sacrificing feeding time or food-quality. We evaluated moose response to roads in areas with and without established wolf packs, testing if wolves add to the landscape of fear. Our study area is in Norway and Sweden where we collected browse data from 2008 to 2018 from fixed quadrats, in areas with and without wolves. We measured moose browsing pressure on important forage species. We will apply mixed effects models to describe the effect of roads on moose browsing in areas with and without wolves using predictor variables wolf presence/absence, road type, distance to nearest road, and distance to forest edge. We will discuss implications to management.

• Eriksen, Ane; Wabakken, Petter; Maartmann, Erling & Zimmermann, Barbara (2017). Do brown bears plan for breakfast? Den site selection for easy access to ungulates in spring. Full text in Research Archive
• Wabakken, Petter; Eriksen, Ane; Maartmann, Erling; Zimmermann, Barbara; Flagstad, Øystein & Liberg, Olof [Show all 8 contributors for this article] (2017). Wolf depredation on livestock, semi-domestic reindeer and domestic dogs in Norway. Full text in Research Archive
• Zimmermann, Barbara; Wabakken, Petter; Eriksen, Ane; Sand, Håkan & Wikenros, Camilla (2017). Who is outside the door? Wolf movement in relation to human settlements. Full text in Research Archive
• Fuchs, Boris; Sørheim, Kristin; Chincarini, Mattheo; Bjarnason, Asgeir; Brunberg, Emma & Stubsjøen, Solveig S. [Show all 11 contributors for this article] (2017). Heart Rate Logging In Free-Ranging Mammals. Full text in Research Archive
• Zimmermann, Barbara (2017). Nytt fra rovviltforskningen på Evenstad. Full text in Research Archive
• Zimmermann, Barbara (2017). Glimt fra ulveforskningen. Full text in Research Archive
• Zimmermann, Barbara & Wabakken, Petter (2017). Ulv og nærhet til mennesker: Slettåsulvenes forflytningsmønster i forhold til hus. Full text in Research Archive
• Zimmermann, Barbara (2017). Big teeth, big wounds: Large carnivores in Scandinavia. Full text in Research Archive
• Zimmermann, Barbara (2017). Ulvene rundt Deset. Full text in Research Archive
• Zimmermann, Barbara (2017). Veiledning til utstillingen Dyr du skulle ønske ikke fantes. Full text in Research Archive
• Zimmermann, Barbara (2017). Hva er forskning? Full text in Research Archive
• Zimmermann, Barbara (2017). Glimt fra ulveforskningen. Full text in Research Archive
• Zimmermann, Barbara (2017). Et ulveår. Full text in Research Archive
• Zimmermann, Barbara (2017). Wolves and people: movement pattern of the Slettås pack in relation to houses. Full text in Research Archive
• Zimmermann, Barbara; Tofastrud, Morten & Vogler, Thomas (2017). Storfe på skogsbeite - Sommeren 2016. Full text in Research Archive
• Tofastrud, Morten; Hegnes, Hilde; Hessle, Anna & Zimmermann, Barbara (2017). Activity and weight gain of free-ranging beef cattle in south-boreal forests of Norway. Full text in Research Archive
• Zimmermann, Barbara (2017). Glimt fra ulveforskningen. Full text in Research Archive
• Zimmermann, Barbara (2017). I ulvens spor. Rovdyrviten. ISSN 1894-1664. p. 30–31. Full text in Research Archive
• Tofastrud, Morten; Hegnes, Hilde; Hessle, Anna & Zimmermann, Barbara (2017). Activity and weight gain of free-ranging cattle in south boreal forests of Norway. Full text in Research Archive
• Tofastrud, Morten & Zimmermann, Barbara (2017). Storfe på skogsbeite. Full text in Research Archive
• Tofastrud, Morten & Zimmermann, Barbara (2017). Habitatvalg og beiteatferd hos kjøttfe i boreal skog. TYRmagasinet. ISSN 1890-8462. p. 8–8. Full text in Research Archive
• Zimmermann, Barbara (2016). Large carnivores of Scandinavia: Status, conflicts, management. Full text in Research Archive
• Zimmermann, Barbara (2016). Elg og rovdyr. Full text in Research Archive
• Sand, Håkan; Mallwitz, Daniel & Zimmermann, Barbara (2016). Älgars flyktbeteende vid jakt med löshund inom och utanför vargområden. Fakta Skog. ISSN 1400-7789. 1, p. 1–4. Full text in Research Archive
• Zimmermann, Barbara (2016). Elgforvaltning i ulverevir. Full text in Research Archive
• Zimmermann, Barbara; Tofastrud, Morten; Vogler, Thomas; Bakmann, Ole & Skogholm, Bjørn (2016). Cattle in the woods: Sustainable beef cattle production on forest pastures. Full text in Research Archive
• Eckhoff, Rikke & Zimmermann, Barbara (2016). Hvorfor er vi så redde for ulven? [Radio]. NRK P2 Ekko. Full text in Research Archive
• Zimmermann, Barbara (2016). Elgforvaltning i ulverevir. Full text in Research Archive
• Eckhoff, Rikke; Tønnessen, Morten; Skogen, Ketil & Zimmermann, Barbara (2016). Hvorfor er vi så redde for ulven? [Radio]. "Ekko" på NRK P2. Full text in Research Archive
• Tofastrud, Morten & Zimmermann, Barbara (2016). Grazing of free-ranging cattle in south-boreal forests of south eastern Norway. Full text in Research Archive
• Zimmermann, Barbara (2015). Utmarksfag på Evenstad. Full text in Research Archive
• Zimmermann, Barbara (2015). Elgforvaltning i ulverevir. Full text in Research Archive
• Zimmermann, Barbara (2015). Large carnivores of Scandinavia: Status, ecology, conflicts. Full text in Research Archive
• Zimmermann, Barbara (2015). Love it and hate it: Scale-dependent ambivalent behaviour of wolves to roads. Full text in Research Archive
• Zimmermann, Barbara (2015). Etablering av hjort i tradisjonelle elgområder. Full text in Research Archive
• Zimmermann, Barbara (2015). Julussa-ulvene: arealbruk og predasjon. Full text in Research Archive
• Meisingset, Erling L.; Rivrud, Inger Maren; Loe, Leif Egil; Zimmermann, Barbara; Brekkum, Øystein & Mysterud, Atle (2015). Hjortens områdebruk om høsten. Hjorteviltet. ISSN 1502-3729. 25, p. 101–103. Full text in Research Archive
• Zimmermann, Barbara (2015). Ulven tar fleire elg enn han treng. [Journal]. Nationen. Full text in Research Archive
• Zimmermann, Barbara & Jakobsen, Siw Ellen (2015). Ulven både elsker og hater skogsbilveiene. [Journal]. Nationen. Full text in Research Archive
• Zimmermann, Barbara (2015). Skogsforbud skaper mer frykt - mener forsker. [Radio]. NRK Østlandssendingen. Full text in Research Archive
• Zimmermann, Barbara; Lien, Ole Mattis; Wedul, Sari Jemiina; Johnsen, Kaja & Østerhus, Benedicte (2015). Hind 359: En hjort med sans for vann og farlige veier. Hjorteviltet. ISSN 1502-3729. 25, p. 80–82. Full text in Research Archive
• Zimmermann, Barbara (2015). Gråbein eter beveren. [TV]. NRK Østlandssendingen. Full text in Research Archive
• Zimmermann, Barbara; Sand, Håkan; Wabakken, Petter; Wikenros, Camilla & Liberg, Olof (2015). Impact of wolf predation on the highly managed Scandinavian moose population. Full text in Research Archive Show summary

  To understand the impact of predation is essential when managing game species. The Scandinavian Wolf Research Project (SKANDULV) estimated winter kill rates by searching GPS-positions of GPS-collared wolves for carcass remains, and derived winter ungulate densities by means of fecal pellet counts. Moose (Alces alces) made up 70% of all prey items found, or 95% of all killed biomass. Wolves killed on average 0.061 moose per wolf per day (range 0.012 – 0.135), and 73% of all moose carcasses were calves. The kill rates were positively related to moose density and negatively to wolf pack size and were best described by a predator-dependent functional response model. Small wolf packs (2 – 4 wolves) acquired more than three times as much biomass as required to cover the energetic needs of the pack, while large packs (7 – 9 wolves) in territories with low to intermediate prey abundance seemed to be food-limited during winter. Predation rates, i.e. the proportion of the moose population killed by predators during winter, were not related to wolf pack size or kill rates, but were negatively correlated with moose abundance or the moose-to-wolf ratio within the wolf territory. Wolves did not regulate, but rather limited, the highly managed moose population in Scandinavia, with wolf-induced mortality being mainly additive to other sources of mortality. With their strong preference for calves, wolves have a smaller impact on moose population dynamics than hunters who harvest a higher proportion of productive animals.

• Kristiansen, Rolf Håkon & Zimmermann, Barbara (2015). Alperose i ulveland. [Journal]. Jakt og Fiske. Full text in Research Archive
• Zimmermann, Barbara & Harstad, Lisbet (2015). Vil beskytte elgkua under jakta. [Journal]. Nationen. Full text in Research Archive
• Zimmermann, Barbara (2015). Ulvens atferd i et menneske-dominert landskap. Full text in Research Archive
• Zimmermann, Barbara (2015). Ulvens uttak av elg. Full text in Research Archive
• Zimmermann, Barbara & Jakobsen, Siw Ellen (2015). Ulven både elsker og frykter skogsbilveiene. [Internet]. forskning.no. Full text in Research Archive
• Zimmermann, Barbara (2015). Elgjakt i ulverevir. Full text in Research Archive
• Zimmermann, Barbara & Jakobsen, Siw Ellen (2015). Ulven dreper flere elg enn den trenger. [Internet]. forskning.no. Full text in Research Archive
• Zimmermann, Barbara (2015). Elgforvaltning i ulverevir. Full text in Research Archive
• Zimmermann, Barbara (2015). Julussaflokken. Full text in Research Archive
• Skogen, Ketil; Odden, John; Linnell, John Durrus; Krange, Olve; Kaltenborn, Bjørn Petter & Støen, Ole-Gunnar [Show all 9 contributors for this article] (2014). Rovdyrsonene fungerer ikke. Full text in Research Archive
• Skogen, Ketil; Odden, John; Linnell, John Durrus; Krange, Olve; Kaltenborn, Bjørn Petter & Støen, Ole-Gunnar [Show all 9 contributors for this article] (2014). Rovdyrsonen fungerer ikke. Full text in Research Archive
• Skogen, Ketil; Odden, John; Støen, Ole Gunnar; Wabakken, Petter; Linnell, John Durrus & Zimmermann, Barbara [Show all 9 contributors for this article] (2014). Rovdyrsonene fungere ikke. Østlendingen. Full text in Research Archive
• Wabakken, Petter & Zimmermann, Barbara (2014). Fem myter om ulv. [Journal]. Aftenposten. Full text in Research Archive
• Zimmermann, Barbara (2014). Wolf predation in Scandinavia. Full text in Research Archive
• Zimmermann, Barbara; Sand, Håkan; Wabakken, Petter; Wikenros, Camilla & Liberg, Olof (2014). Wolf predation in Scandinavia: Prey selection, functional response, energetics and predation rates. Full text in Research Archive
• Zimmermann, Barbara; Cederström, Madelen & Kjæstad, Reidar (2014). Frykter for elgjakta i Trysil. [Radio]. NRK. Full text in Research Archive
• Wabakken, Petter; Zimmermann, Barbara & Nærland, Mina Hauge (2014). Fem myter om ulv. [Journal]. Aftenposten. Full text in Research Archive
• Zimmermann, Barbara & Eide, Stian (2014). Spiser elg for millioner. [Journal]. Nationen. Full text in Research Archive
• Zimmermann, Barbara & Sveen, Knut (2014). Spår slutten på elgjakta. [Internet]. NRK. Full text in Research Archive
• Zimmermann, Barbara (2014). Large carnivore predation: Wildlife versus livestock. Full text in Research Archive
• Zimmermann, Barbara (2014). Kåseri om rovdyr. Full text in Research Archive
• Zimmermann, Barbara (2013). Large carnivores in Norway. Full text in Research Archive
• Zimmermann, Barbara; Wabakken, Petter; Taylor, Lindsey; Sand, Håkan & Liberg, Olof (2013). Behavioral response of wolves to roads: Scale-dependent ambivalence. Full text in Research Archive
• Zimmermann, Barbara (2013). Hjort i Østerdalen. Full text in Research Archive
• Zimmermann, Barbara (2013). Hjorten har kommet for godt. Full text in Research Archive
• Kaja, Johnsen; Zimmermann, Barbara; Milner, Jocelyn Margarey & Wedul, Sari Jemiina (2013). Forstyrrelseskonkurranse: Vil hjorten utkonkurrere elgen på fôringsstajonene? Hjorteviltet. ISSN 1502-3729. p. 40–42. Full text in Research Archive
• Zimmermann, Barbara; Johnsen, Kaja & Wedul, Sari Jemiina (2013). Menneskeskapte matressurser styrer områdebruken til hjorten. Hjorteviltet. ISSN 1502-3729. 2013, p. 37–39. Full text in Research Archive
• Mysterud, Atle; Meisingset, Erling L.; Loe, Leif Egil; Bischof, Richard; Zimmermann, Barbara & Veiberg, Vebjørn (2012). Hjortens trekkmønster. Hjorteviltet. ISSN 1502-3729. 22, p. 38–45. Full text in Research Archive
• Solheim, Kristin; Isaksen, Runo Andre & Zimmermann, Barbara (2012). Utdanner nordiske naturforvaltere. [Journal]. Nordplus 2008-2011. Full text in Research Archive
• Isaksen, Runo Andre & Zimmermann, Barbara (2012). Utdanner nordiske naturforvaltere. [Internet]. www.siu.no. Full text in Research Archive
• Zimmermann, Barbara (2012). Om den store stygge ulven og den søte stygge storken. Full text in Research Archive
• Myklebust, Kjetil & Zimmermann, Barbara (2012). Birkebeinerritt eller naturvern? [Internet]. www.nordplusonline.org. Full text in Research Archive
• Zimmermann, Barbara; Taylor, Lindsey; Wabakken, Petter; Sand, Håkan & Liberg, Olof (2012). Love it AND hate it:Scale-dependent ambivalent behavioural responses of wolves to roads. Full text in Research Archive
• Isaksen, Runo Andre & Zimmermann, Barbara (2012). Nordiske naturforvaltere etterlyses. [Internet]. www.forskning.no. Full text in Research Archive
• Nilsen, Erlend Birkeland; Gervasi, Vincenzo; Linnell, John D. C.; Odden, John; Pedersen, Hans-Christian & Wabakken, Petter [Show all 8 contributors for this article] (2011). Effekten av predasjon på hjorteviltbestander. Hjorteviltet. ISSN 1502-3729. p. 91–93. Full text in Research Archive
• Eggum, Eli Tangen & Zimmermann, Barbara (2011). Hjorten kommer. [Journal]. Landbruksbladet for Kongsvinger. Full text in Research Archive
• Zimmermann, Barbara (2011). Prosjekt Hjort i Hedmark. Full text in Research Archive
• Zimmermann, Barbara; Wabakken, Petter; Sand, Håkan; Strømseth, Thomas Holm; Liberg, Olof & Pedersen, Hans-Christian (2011). Wolves in Scandinavia: Monitoring, research and management. Full text in Research Archive
• Zimmermann, Barbara (2011). Hjort i Hedmark: arealbruk og trekk. Full text in Research Archive
• Zimmermann, Barbara (2011). Ulveforskning på Evenstad. Full text in Research Archive
• Zimmermann, Barbara (2011). Stanges unike hjortestamme. [Journal]. Hamar Arbeiderblad (HA). Full text in Research Archive
• Zimmermann, Barbara (2011). Ulvens predasjon på elg. Full text in Research Archive
• Zimmermann, Barbara (2010). GPS-merking av hjort på Haugalandet. Full text in Research Archive
• Zimmermann, Barbara (2010). Wolf-moose research in Scandinavia. Full text in Research Archive
• Loe, Leif Egil; Mysterud, Atle; Meisingset, Erling L.; Veiberg, Vebjørn & Zimmermann, Barbara (2010). Arealbruk hos hjort, med nye data fra Buskerud. Full text in Research Archive
• Gangås, Kristin Evensen; Andreassen, Harry P.; Zimmermann, Barbara & Wabakken, Petter (2010). Predicting wolf distribution by means of historical data. Full text in Research Archive Show summary

  One big challenge in management of human-wildlife conflicts is the lack of predictability of where large carnivores will establish in the future. Without such forecasts it is impossible to know where to introduce preventive actions to decrease conflicts. In Scandinavia, we have longitudinal monitoring data on the establishment of the Scandinavian wolf population since late 1970s. We will develop models based on Kernel density distributions to predict the future establishment of wolf populations. In a second stage we will combine this with spatial distributions of economic and social factors to improve spatial management of large carnivores.

• Wikenros, Camilla; Zimmermann, Barbara; Sand, Håkan; Månsson, Johan; Maartmann, Erling & Eriksen, Ane [Show all 7 contributors for this article] (2010). Effect of predation on hunting quotas, hunter observations and moose harvest in two countries with an increasing wolf population during 22 years. Full text in Research Archive
• Zimmermann, Barbara; Willebrand, Tomas & Torkar, Gregor (2010). An education model for future managers to improve sustainable conservation efforts. Full text in Research Archive Show summary

  Sustainable conservation is put into action by different stakeholders and managers at the local, regional, national and international level. A multidisciplinary framework in the education of future managers is therefore crucial for the success of conservation projects. We present an educational model that does not only include the ecological in- depth knowledge, but also the human dimension, the ability to cooperate, and the international perspective. In a Nordic-Baltic joint effort financed by Nordplus, we organized an intensive course with students from ten universities. Mixed student groups were studying a conflict regarding natural resources by means of literature research and qualitative interviews of stakeholders. Students had a high learning outcome.

• Zimmermann, Barbara; Sand, Håkan; Wabakken, Petter & Andreassen, Harry P. (2010). Functional response in Scandinavian wolves: implications for sustainable management of a conflict species. Full text in Research Archive Show summary

  We estimated the functional response of 12 wolf packs interacting with moose and roe deer as prey in 16 winter studies in Scandinavia. Wolves showed a strong predator dependent functional response. While large packs killed just about the amount of prey required to cover the daily requirements, smaller packs had higher per capita kill rates, and pairs exceeded by far their daily requirements. If wolves are controlled to keep the impact on moose below a certain threshold, this may be more successful by limiting the establishment of new packs rather than removing individuals from already established packs.

• Zimmermann, Barbara (2010). Wolves in Scandinavia. Full text in Research Archive
• Zimmermann, Barbara; Sand, Håkan; Wabakken, Petter; Strømseth, Thomas Holm & Andreassen, Harry P. (2010). Functional response of wolves in Scandinavia: Implications for sustainable management of a conflict species. Full text in Research Archive
• Zimmermann, Barbara (2010). Hjort i Hedmark, GPS på hjort. Full text in Research Archive
• Zimmermann, Barbara (2009). Ulv i Fulufjellet. Full text in Research Archive
• Zimmermann, Barbara; Meisingset, Erling L.; Loe, Leif Egil & Mysterud, Atle (2009). Hjortens områdebruk: Hvem blir igjen og hvem trekker? Full text in Research Archive
• Zimmermann, Barbara (2008). Wolf predation in Scandinavia. Full text in Research Archive
• Zimmermann, Barbara (2008). Wolf and moose in Scandinavia. Full text in Research Archive
• Zimmermann, Barbara (2008). Store rovdyr i Skandinavia. Full text in Research Archive
• Zimmermann, Barbara; Wabakken, Petter & Strømseth, Thomas Holm (2008). Status for ulv og ulveovervåking. Full text in Research Archive
• Zimmermann, Barbara (2008). Ulv og elg i Skandinavia. Full text in Research Archive
• Zimmermann, Barbara (2008). GPS merkeprosjekter på hjort i Norge. Full text in Research Archive
• Zimmermann, Barbara (2008). Hjort på Haugalandet - hva har vi lært fra merkeprosjektet? Full text in Research Archive
• Gundersen, Hege; Zimmermann, Barbara; Milner, Jos M.; Fjone, Gunnar & Andreassen, Harry P. (2007). Spatial economic analyses in moose management. Full text in Research Archive Show summary

  The costs and benefits related to moose and moose hunting in Norway varies considerable depending on whether a landowner?s property lies in a traditional wintering area or summer area for moose. During winter the moose can cause major damage to young pine forest, whereas during late summer/autumn the moose represents great values due to hunting related income. Typically, moose in central Norway stay in lower elevated areas during winter, but migrates to higher altitudes when calving season starts. This seasonal migration pattern is induced by two important climatic factors; snow depth and temperature. By the use of kernel density functions based on moose telemetry data we have modelled spatial and temporal changes in moose densities for two different seasons through several years. The utilisation distributions (UD) are matched with GIS data on climate and altitude in a cell-by-cell fashion explained by Marzluff et al. (2004). An economic moose management model is then presented, based on predicted spatiotemporal moose density maps and the actual landowner configuration in the area, suggesting a fair allocation of income and costs related to moose and moose hunting in Norway.

• Gundersen, Hege; Zimmermann, Barbara; Storaas, Torstein & Nicolaysen, Knut (2007). Fire elger tog-drept på én natt. Full text in Research Archive
• Zimmermann, Barbara; Wabakken, Petter; Strømseth, Thomas Holm; Pedersen, Hans Christian; Sand, Håkan & Liberg, Olof (2007). Ulv og elg på låven. Full text in Research Archive
• Sand, Håkan; Wabakken, Petter; Zimmermann, Barbara; Pedersen, Hans Christian; Liberg, Olof & Wikenros, Camilla (2007). Predation ecology of wolves in Scandinavia. Full text in Research Archive
• Zimmermann, Barbara; Sand, Håkan; Wabakken, Petter; Pedersen, Hans Christian & Liberg, Olof (2007). Estimation of wolf kill rates with GPS-technology and GIS cluster analysis. Full text in Research Archive Show summary

  To estimate wolf (Canis lupus) kill rates from fine-scale movement patterns, we followed adult wolves in 3 territories of the Scandinavian wolf population using GPS (Global Positioning System) during the winters of 2001 to 2003. The resulting 6 datasets of 62 to 84 study days gave a total of 8,747 hourly GPS positions. We visited clusters of positions in the field on average 8.8 days after positioning and found moose (Alces alces) killed by wolves during the study period on 74 (8%) of the 953 clusters. The number of positions and visits to a cluster and the proportion of afternoon positions were significant fixed effects in mixed logistic regression models predicting the probability of a cluster containing a wolf-killed moose. The models, however, displayed a poor predictability and were not a suitable tool for estimating kill rates from positioning data alone. The proportion of wolf-killed ungulates to be included in clusters depended strongly on both number of positions per day and radius to define a set of spatially aggregated positions as a cluster. A higher proportion (78%) of killed prey were in clusters based on night time positions than in those based on daytime positions (41%). Simulation of aerial search for killed moose during daylight hours resulted in a serious underestimation (>60%) as compared to the number of wolf-killed moose found when using cluster analysis. We discuss proximate and ultimate factors, i.e. human disturbance, access to prey, social organization, intra-guild dominance, and litter size, as potential causes of the observed high temporal and spatial variation in prey handling.

• Gundersen, Hege; Zimmermann, Barbara; Milner, Jocelyn Margarey; Fjone, Gunnar & Andreassen, Harry Peter (2007). Spatial economic analyses in moose management. Full text in Research Archive Show summary

  The costs and benefits related to moose and moose hunting in Norway varies considerable depending on whether a landowner?s property lies in a traditional wintering area or summer area for moose. During winter the moose can cause major damage to young pine forest, whereas during late summer/autumn the moose represents great values due to hunting related income. Typically, moose in central Norway stay in lower elevated areas during winter, but migrates to higher altitudes when calving season starts. This seasonal migration pattern is induced by two important climatic factors; snow depth and temperature. By the use of kernel density functions based on moose telemetry data we have modelled spatial and temporal changes in moose densities for two different seasons through several years. The utilisation distributions (UD) are matched with GIS data on climate and altitude in a cell-by-cell fashion explained by Marzluff et al. (2004). An economic moose management model is then presented, based on predicted spatiotemporal moose density maps and the actual landowner configuration in the area, suggesting a fair allocation of income and costs related to moose and moose hunting in Norway.

• Zimmermann, Barbara; Gundersen, Hege; Hjeltnes, Arne; Sageie, Jostein & Rosef, Olav (2007). Modelling resource utilisation of temporally irregular and spatially autocorrelated GPS data of red deer. Full text in Research Archive Show summary

  The application of resource selection functions (RSF) for large datasets of GPS-positions is limited by the relatively high proportion of used grid cells compared to the available area. Resource utilisation functions (RUF) assume that the degree of use of areas without relocations follows a kernel density distribution of the relocations (Millspaugh et al. 2006). This assumption though is often violated by the fact that most relocation techniques are discrete in time and therefore display only a narrow time window of resource use. We have modified the RUF method by relating kernel density values to available resources at the site of relocations only. Contrary to grid cells, relocations are not systematically distributed in space and often not systematically sampled in time, and we accounted for temporal and spatial autocorrelation by applying the time kernel method (Katajisto and Moilanen 2006) and subsampling relocations with a minimum nearest neighbour distance defined by the relocation error and the map pixel size of the resources. We used generalized linear mixed models (GLMM) with a spatial covariance matrix and different combinations of random intercept and coefficients (Gillies et al. 2006) to describe resource use. Different resource variables were derived from vegetation maps based on Landsat images and from a terrain model.

• Fuchs, Boris; Thiel, Alexandra; Græsli, Anne Randi; Lund, Signe Helene; Evensen, Mari Tangen & Buhler, Kayla Joy [Show all 11 contributors for this article] (2026). Hvorfor er elgkalvene mindre enn før? Universitetet i Innlandet. ISSN 9788283806298. Full text in Research Archive Show summary

  Slaktevektene hos elgkalver (Alces alces) i Norge har gått betydelig ned de siste tiårene, særlig i sørlige og kystnære områder. Denne rapporten oppdaterer vår analyse fra 2024 med et utvidet datasett fra 13 fylker (Del 1–2), og undersøker klimatiske, landskapsmessige, sykdoms- og bestandsrelaterte faktorer. Totalt inngår 171 186 slaktevekter fra perioden 1985–2024. Ved bruk av en strukturell ligningsmodell (SEM) analyserte vi hvordan klima, flåttforekomst, tømmeravvirkning (som proxy for beitetilgang), elgtetthet, rekruttering og maternelle forhold påvirker kalvenes slaktevekt. Alle fylker viste nedgang, størst i Agder (−39%) og minst i Finnmark (−2%). Dagens vekter er lavest i sør (40–45 kg) og høyest i nord (>60 kg). Lavere vekter var knyttet til varmere somre, høyere flåttbelastning og dårligere kondisjon hos kyr. Kyrnes kondisjon hadde sterkest positiv effekt. Resultatene viser at nedgangen skyldes samspill mellom klima, flått og mordyrkondisjon. Forvaltningen kan bidra ved å holde bestanden under bæreevnen og sikre gode beite- og leveområder.

• Spedener, Mélanie; Austrheim, Gunnar; Goudier, Alexandre; Hauer, Josh; Helbert, Juliette & Henriksen, Marie Vestergaard [Show all 16 contributors for this article] (2025). Husdyr på skogsbeite: Påvirkninger på granforyngelse og plante-pollinator-samfunn. Universitetet i Innlandet. ISSN 9788283805871. Full text in Research Archive Show summary

  Diskusjonen om husdyr på utmarksbeite i skog er sammensatt og går blant annet ut på hvordan det kan kombineres med skogdrift, rovviltforvaltning, og bevaring av biologisk mangfold. I dette forskningsprosjektet har vi undersøkt hvordan husdyr på sommerbeite i skog innenfor eller nær ulvesonen påvirker skogbruk og artsmangfold. Nærmere bestemt har vi studert effektene av sau innenfor rovviltavvisende gjerder og av frittgående storfe på granforyngelse og plante-pollinator samfunn i boreal produksjonsskog. Vi undersøkte hvorvidt beitedyrene gjorde skade på unge grantrær gjennom beiting og tråkk og hvorvidt de bidro til ungskogspleie ved å fjerne konkurrerende vegetasjon. I tillegg undersøkte vi hvordan beitedyr påvirket planter i feltsjiktet, blomsterressurser og pollinerende insekter. Hverken storfe eller sau førte til økte beite- og tråkkeskader på små grantrær. Både storfe og sau bidro til ungskogspleie ved å beite på løvtrær og ved å redusere høyden av feltsjiktvegetasjonen. Storfe førte til økt dødelighet hos gran, om så i svært begrenset omfang, mens sau førte til økt høydetilvekst. Både storfe og sau påvirket feltsjiktvegetasjonen, men storfe i større grad enn sau. Sau førte til redusert høyde av graminider, mens storfe førte til redusert høyde av graminider, lyng og urter, i tillegg til redusert mengde av graminider. Både storfe og sau påvirket blomsterressursene, og storfe hadde igjen større effekt enn sau. Mens sau førte til en nedgang i artsrikdom, førte storfe til nedgang i både artsrikdom og mengde av blomster. Både storfe og sau påvirket pollinerende insekter, og her førte begge til en nedgang i mengden av disse. Funnene våre tilsier at både sau innenfor rovviltavvisende gjerder og frittgående storfe godt kan kombineres med dagens skogbruk som har som hovedmål å produsere grantømmer ved bruk av flatehogst – i hvert fall når husdyrtetthetene er like lave som i våre studieområder. Vi var overrasket over at husdyrene hadde negative effekter på plante-pollinator samfunn. Dette er bekymringsfullt og bør tas hensyn til, særlig siden vi nesten utelukkende fant ville pollinatorer i studieområdene våre, som er av stor interesse i forhold til naturvern.

• Spedener, Mélanie; Zimmermann, Barbara; Mathisen, Karen Marie; Tofastrud, Morten; Austrheim, Gunnar & Henriksen, Marie Vestergaard [Show all 7 contributors for this article] (2025). The role of summer grazing beef cattle in boreal production forest. Universitet i Innlandet / University of Inland Norway. ISSN 9788283805543. Full text in Research Archive Show summary

  Sammendrag på norsk I Norge beiter kjøttfe i store områder av boreal produksjonsskog preget av flatehogst på sommeren (mai-september). Vi studerte først mat- og habitatvalg av disse kyrne (Artikkel I– II), og deretter effektene av storfe på flora og fauna (Artikkel III-V). Datainnsamlingen foregikk i Sørost-Norge i 2015-2017 (Furnes/Vang og Stange/Romedal) og 2021-2023 (Steinvik og Deset). Vi studerte kyrnes ressursvalg ved å klassifisere deres adferd ved hjelp av GPS og akselerasjonsdata, ved å hente inn (fra kart) og måle (i felt) habitatvariabler, ved å samle inn møkkprøver til mikrohistologiske analyser og ved å modellere ressursseleksjonsfunksjoner. Vi fokuserte på unge granplantefelt for å studere effektene av kjøttfe på flora og fauna, siden kyrene selekterer for denne skogstypen. Dessuten har små grantrær høy økonomisk verdi og unge granplantefelt er rikere i blomster og pollinatorer enn det resterende skoglandskapet. På 24 unge granplantefelt satt vi opp parede prøveflater (20x20 m hver), hvorav en omgitt av et gjerde. Vi så på unge trær, vegetasjonen i feltsjiktet og blomsterbesøkende insekter. Siden halvparten av disse granplantefeltene lå innenfor, og den andre halvparten utenfor beiteområdene, kunne vi skille effektene av storfe fra effektene av hjortedyr, som lever vilt i disse skogene. Interaksjoner mellom storfe og hjortedyr studerte vi ved å sette opp viltkamera på de samme granplantefelt og ved å gjennomføre møkktellinger langs et rutemønster i ett av beiteområdene. Kyrne hadde en gressrik diett og selekterte for gressrike habitater, både på stor og på liten skala (Artikkel I). Storfe selekterte for forskjellige habitatvariabler (liten skala) avhengig av adferden: Når de beitet, selekterte de for gressrikt habitat, og når de hvilte, selekterte de for gressrikt habitat med lite helling og høy kronedekning (Artikkel II). Storfe førte til bittelitt høyere dødelighet av unge grantrær, men ikke til høyere risiko for tråkk- og beiteskader (Artikkel III). Storfe fjernet vegetasjon som konkurrerte med unge grantrær, det vil si unge løvtrær og vegetasjon i feltsjiktet (Artikkel III). Storfe påvirket plante-pollinatorsamfunnet på en annen måte enn hjortevilt: Utgjerding av klovdyr utenfor beiteområde (hjortedyr) førte til lavere abundans av blomster, mens utgjerding av klovdyr innenfor beiteområde (hjortedyr og storfe) førte til lavere abundans av blomster og lavere abundans av blomsterbesøkende insekter (Artikkel IV). Elg brukte andre habitattyper enn storfe (Artikkel V). Elgen sitt bruk av unge granplantefelt avtok med økende bruk av storfe (Artikkel V). Mulige beiteinnskrenkende tiltak, samt bevaring av artsmangfoldet i boreal produksjonsskog ble drøftet, og anbefalinger for videre forskning ble gitt.

• Ausilio, Giorgia; Zimmermann, Barbara; Sand, Håkan; Wikenros, Camilla; Eriksen, Ane & Wabakken, Petter [Show all 12 contributors for this article] (2023). Gränsöverskridande älgförvaltning - Ekologiska förutsättningar. Universitetet i Innlandet. ISSN 9788283804089. Full text in Research Archive Show summary

  Gjensidige påvirkninger mellom rovdyr og byttedyr formes av flere ulike faktorer inklusive type landskap de lever i og rovdyrenes jaktmetode. I landskap påvirket av mennesker kan disse interaksjonene endres av aktiviteter som jakt, skogbruk og andre typer av arealbruk. I Skandinavia har jakt blitt den største dødsårsaken for elg (Alces alces) i det siste århundret, men med tilbakekomsten av store rovdyr slik som bjørn (Ursus arctos) og ulv (Canis lupus) i de seneste årene blir dødsrisikoen nå fordelt på flere kilder. Målet med denne studien var å 1) beskrive habitatet på de plassene der elg ble skutt av jegere i jaktsesongen, og der elg ble drept av ulv både i og etter jaktsesongen, for å finne ut om risikoen for å dø av de to årsakene var koblet til spesifikke habitatstyper; 2) undersøke om habitatvalget til GPS-merket elg påvirkes av fordelingen av jakt- og predasjonsrisiko i tid og rom; og 3) beregne dødeligheten hos elgkalver sommer- og vinterstid i forhold til risiko fra jakt, store rovdyr (ulv og bjørn) og andre miljøvariabler. Vi fant at plassene der elg ble skutt skilte seg fra plassene der elg ble drept av ulv på mange ulike måter som var koblet til habitat i form av alder og type skog, elgtetthet og nærhet til bebyggelse. I neste steg koblet vi disse resultatene til habitatvalget til GPS-merket elg. I jaktsesongen unngikk elgene habitat med høy jaktrisiko på dagtid, mens deres habitatvalg nattestid ikke var koblet til risiko for å bli skutt. Etter jaktsesongen fantes det ingen kobling mellom elgenes habitatvalg og jaktrisiko, hverken på dagtid eller nattestid. Derimot valgte GPS-elgene habitat med høyere risiko for ulvepredasjon både på dagtid og nattestid i og etter jaktsesongen. Dødeligheten hos elgkalver sommerstid varierte signifikant mellom de tre ulike studieårene, men var ikke relatert til risiko for ulvepredasjon, bjørnetetthet, eller klima- eller miljøvariabler. I vinterperioden var kalvedødeligheten positivt koblet til snødybde og andelen hogstflater og ungskog innenfor ulverevir, samt til habitat med økt jaktrisiko. Oppsummert viser disse resultatene at selv om risikoene fra både jakt og ulvepredasjon varierte i tid og rom, så responderte elgene atferdsmessig hovedsakelig på den sterkere og mer forutsigbare dødelighetskilden; jakt. Disse resultatene er sannsynligvis en effekt av at jakt er den dominerende dødsårsaken, som funksjonelt har erstattet predasjon fra ulv og bjørn på elg i løpet av det siste århundret i Skandinavia.

• Wikenros, Camilla; Zimmermann, Barbara; Eriksen, Ane; Esselin, Anders; Falkevik, Maria & Løken, Øivind [Show all 9 contributors for this article] (2023). GRENSEVILT 2 - Slutrapport till Interreg Sverige-Norge. Universitetet i Innlandet. ISSN 9788283804058. Full text in Research Archive Show summary

  Indre Skandinavia kjennetegnes av et sammenhengende barskogområde med store viltbestander (naturarv) som preger regionens skogbruk og jakt (kulturarv). Til tross for at naturressursene er grenseoverskridende, forvaltes og utnyttes de ulikt på hver sin side av administrative grenser. Det fører til interessekonflikter og ugunstige forvaltningsmodeller og forvaltningsenheter som ikke tar hensyn til dyrenes økologi. Vindkraftutbygging har akselerert i Indre Skandinavias barskogområder, uten at man er klar over konsekvensene for viltbestander generelt og på tvers av forvaltningsregioner spesielt. Det er et stort behov for forvaltningsrelevant kunnskap som tilrettelegger for og forbedrer grenseoverskridende samarbeid. GRENSEVILT 2 (2020-2022) er en oppfølging av prosjektet GRENSEVILT 1 (2017-2021). Ny kunnskap og ressursgruppens arbeid førte til et uttrykt ønske fra forvaltningen, grunneiere og interesseorganisasjoner i Sverige og Norge om en konkretisering av samarbeidet samt økt kunnskapsinnhenting som underlag for samarbeidet. GRENSEVILT 2 hadde som overordnet mål å øke samordningen av viltforvaltning på tvers av grensen og i områder under press fra menneskelige inngrep. Projektet har hatt to delmål med fokus på å forstå forvaltningsmessige og biologiske forutsetninger for å oppnå et bedre forvaltningssamarbeid, og å forstå effekter av vindkraftutbygging på vilt og jakt. Prosjektets arbeid mot de to delmålene har hatt to hovedkomponenter: 1) oppbygging av et forskningsbasert, forvaltningsrelevant kunnskapsgrunnlag, og 2) nettverks- og kommunikasjonsarbeid på ulike nivåer. GRENSEVILT 2 følger Interregs prosess til å skape regional merverdi gjennom å bli kjent med hverandre, dele kunnskap, og finne fram til felles, grenseoverskridende løsninger. Prosjektgruppen bestod av forskere og forvaltere fra Norge og Sverige, samt en prosessleder. I GRENSEVILT 2 prosjektet har vi lagt en struktur for regionalt samarbeid på tvers av grensen. Vi har økt forståelsen av forvaltningsmessige, metodiske og økologiske forutsetninger for samarbeid i elgforvaltningen. Gjennom scenariediskusjoner, oppbygging av en felles kartløsning, og ny kunnskap har prosjektet diskutert nye adaptive forvaltningsmodeller som tar hensyn til elgens vandringer på tvers av grensen og ulike interesser knyttet til skog, jakt, elg, ulv og jerv. GRENSEVILT 2 har tatt fram mer kunnskap om effekter av den stadig økende utbygging av vindkraft i Indre Skandinavias felles skogressurser. Forflytningsdata fra elg, ulv og jerv merket med GPS-halsbånd samt elgmøkktellinger ble analysert for å øke forståelsen av hvordan slike tekniske inngrep i skoglandskapet kan påvirke atferden og områdebruken til ville dyr. Vi har også kartlagt jaktuttaket av elg lokalt og jegernes opplevelse av vindkraftutbyggingen. Under hele prosjekttiden har vi hatt et nært samarbeid med ressursgruppen, som var satt sammen av norske og svenske representanter fra forvaltningen, grunneiere, næringslivet, forskningen og interesseorganisasjoner. Denne gruppen hadde en avgjørende rolle i prosjektet med en tett møteplan og fortløpende prosessledelse. Kommunikasjon var helt sentralt for å kunne nå vårt hovedmål om økt samordning av viltforvaltningen og vi har fulgt en kommunikasjonsplan under prosjektets gang. I GRENSEVILT 2 utnyttet vi strukturene som ble bygget i GRENSEVILT 1 (hjemmeside, Facebook-side, kontaktnett for nyhetsbrev, mediakontakter, lokale, regionale og nasjonale fagfora for foredrag) men har også arbeidet med andre kommunikasjonskanaler (kartfortellinger og film). I tillegg har vi forberedt innholdet i den planlagte internasjonale forsknings- og forvaltningskonferansen «Wolves Across Borders» som avholdes i mai 2023.

• Zimmermann, Barbara; Wikenros, Camilla; Eriksen, Ane; Sand, Håkan; Wabakken, Petter & Bernardi, Cecilia Di [Show all 14 contributors for this article] (2023). Vilt i vind - Pilotstudier om vindkraftutbyggingens påvirkninger på elg, ulv, jerv og elgjegere. Universitetet i Innlandet. ISSN 9788283804096. Full text in Research Archive Show summary

  Abstract As part of the green transition, substantial wind power development is currently taking place in Scandinavia. The establishment of wind power plants leads to environmental changes in the form of logging, infrastructure such as turbines, transformers, and power lines, and a new or improved road network. The turbines and rotor blades generate noise and flicker from reflected sunlight and shade, and result in increased mortality in birds. In addition, there may be a risk of ice throw from the turbines. Little is known about wind power development effects on wildlife and hunting in Scandinavia. In the GRENSEVILT 2 project, we had the opportunity to carry out pilot studies on the possible effects of wind power development on moose, wolves, wolverines, and moose hunting, partly due to the construction of Kjølberget Wind Power Plant at the Norwegian side of the national border in northern Finnskogen area during 2019-2021. Moose pellet counts at the Kjølberget Wind Power Plant indicated a possible negative effect of the construction work that took place in the summer of 2021 on the distribution of moose. This pattern coincides with the area use of the two GPS-marked moose cows that both had their summer areas at the power plant. One stayed away from the turbines in 2021 and the other was most often in close proximity to the turbines at night and in late autumn when the construction work was over. As for wolves, in order to assess the extent to which wind turbines can be expected to affect the wolf's habitat use we investigated whether areas with planned wind power development within the range of the Scandinavian wolf population coincide with important wolf habitat. We found that planned turbine locations were on the periphery as well as in the activity centers of the wolves, and that there was greatest overlap between planned turbine locations and the wolf activity centers during the denning period. The overlap was slightly less late in the summer, and least in the winter. In the lack of access to GPS-collared wolverines that overlapped with planned or established wind power plants, we analyzed the elevation use of GPS-collared wolverines in areas without wind power plants. Reproducing females during the denning period preferred higher elevations, and their dens were most often at the highest elevations within their home ranges. However, this selection for higher elevation areas was not reflected in a disproportionate use of areas with more wind, and the average wind speed in areas selected by wolverines was below what is considered sufficient for wind power development. Nevertheless, our preliminary conclusions are that the wolverine as a species, and especially in potential breeding areas, should be highly topical with regard to future assessments of impact of land-based wind power development in the Scandinavian countryside. We conducted a web-based, anonymous survey aimed at a selection of hunters in Värmland and Dalarna counties in Sweden, and five municipalities in the far east of Innlandet county in Norway. A total of 656 hunters responded. The results show that many hunters perceive wind turbines on or adjacent to their own hunting grounds as significantly negative for both the hunting performance and the nature experience associated with moose hunting. A clear majority of those questioned had a negative attitude towards the continued development of wind power in general, and this negative attitude became even more evident when it came to development on or in proximity to their own hunting grounds. Our results also indicate that there is a stronger negative attitude towards wind power development in general among the Norwegian than the Swedish hunters. An analysis of hunting statistics from moose management areas at wind power plants indicated a low number of moose shot in hunting areas with many turbines and in close proximity to wind power plants, especially during the construction phase. As possible reasons, we discuss reduced moose density in close proximity to wind power plants because moose avoid these areas, and/or reduced harvest because hunters choose to hunt less in areas with wind power plants. In large moose management areas, the effect of wind power plants on moose harvest was less pronounced, and other factors, such as density of wolves and bears, had a larger impact. Our pilot studies point to a great need for research to uncover the effects of wind power development on wildlife and hunting. This requires long-term studies with a before-during-after design, including studies on the behavioral and physiological responses of wild animals to wind power development.

• Hellbaum, Paige; Ampe, Natacha; Carlomagno, Ester; Grimsgaard, Aslak; Luksengård, Sigurd & Miltz, Cecilia [Show all 7 contributors for this article] (2023). Elgbeitetakst 2022 i Våler og Åsnes kommuner. Universitetet i Innlandet. ISSN 9788283804027. Full text in Research Archive Show summary

  På oppdrag fra Våler og Åsnes kommuner gjennomførte Høgskolen i Innlandet en beitetakst for elg (Alces alces) i mai og juni 2022. Metoden bestod av en kombinasjon av den norske (Solbraa) og svenske (Äbin) beitetakstmetodikken. Denne nye metoden gjør det enklere å sammenligne resultater fra begge sider av grensen. Prøveflatene ble valgt ut ved å besøke på nytt de bestand som inngikk i en stor beitetakst i 2021 i regi av Interreg‐prosjektet GRENSEVILT, også la vi i tillegg ut prøveflater i bestand som var undersøkt i en beitetakst fra 2018. Beitegraden på furu (Pinus sylvestris) ble beregnet som andelen av furuskuddene på inntil 10 trær per prøveflate som var beitet. Skadegraden ble beregnet som andelen furutrær som var skadd i løpet av de siste 12 månedene med enten toppskuddbeite, knekt stamme eller barkgnag. Dessuten kartla vi akkumulerte beiteskader ved å beskrive skadebildet for de siste årene for hvert av inntil ti furutrær per prøverflate langs en gradient fra uskadd til ødelagt. Gjennomsnittlig beitegrad og skadegrad på furu i hele området lå på 8%. Beitegraden var høyest for furutrær i nedre Flisdalen i Åsnes kommune, mens bestand med høy skadegrad var spredt over hele området. Omtrent 70% av de undersøkte furutrærne hadde ingen eller bare lette akkumulerte skader. Bestand med økte akkumulerte skader var konsentrert til Risberget i Våler kommune, vest for Glomma i Våler, og noe også i nedre Flisdalen i Åsnes. Tettheten av utviklingsdyktige furutrær på furudominerte prøveflater var for 71% av bestandene høyere enn 150 trær/daa og dermed innenfor anbefalt tetthet av furu etter ungskogpleie. Siden 2013/2014 har beitegraden på furu gått ned og er nå bare omtrent en tredjedel av nivået 8‐9 år tidligere. Skadegraden ble for første gang målt i 2021 og var da litt over dobbelt så høy som i 2022. En eksepsjonelt høy skadegrad i 2021 og en sterk reduksjon i 2022 ble også funnet i den offisielle Äbin‐taksten gjennomført av Skogsstyrelsen på de svenske naboarealene. Andelen trær uten eller med lette akkumulerte skader har økt siden forrige takst utført i 2018 med nesten 20 prosentpoeng. Nedgangen i beitegraden, skadegraden og akkumulerte skader på furu tyder på at vinterbestanden av elg er lavere enn tidligere, til tross for at elgforvaltningen ble lagt opp til å tillate en økning i bestanden. Fordi store deler av Våler og Åsnes kommune er foretrukne vinterområder for elg, er avkytningsstrategier i naboområdene avgjørende for hvor mye elg som konsentrerer seg om vinteren i de to kommunene. Spesielt på svensk side har avskytingen økt for å få ned beiteskader på furu. Ved beregning av kvoter bør det også tas hensyn til ulvens (Canis lupus) uttak av elg.

• Liberg, Olof; Sand, Håkan; Wikenros, Camilla; Åkesson, Mikael; Zimmermann, Barbara & Wabakken, Petter (2022). Spatial dynamics in the Scandinavian wolf population – effects of increased population size and culling. Sveriges Lantbruksuniversitet. Full text in Research Archive
• Wikenros, Camilla; Zimmermann, Barbara; Sand, Håkan; Eriksen, Ane; Wabakken, Petter & Bernardi, Cecilia Di (2022). Avskjutning av älg över tid och rum – effekter av rovdjur och skogsbruk. Sveriges Lantbruksuniversitet. ISSN 9789157699428. Full text in Research Archive
• Sand, Håkan; Zimmermann, Barbara; Wikenros, Camilla & Wabakken, Petter (2022). Predationsstudier på varg inom projekt GRENSEVILT – en jämförelse mellan nya och tidigare studier. Sveriges Lantbruksuniversitet. ISSN 9789157699435. Full text in Research Archive
• Zimmermann, Barbara; Wikenros, Camilla; Eriksen, Ane; Aronsson, Malin; Ausilio, Giorgia & Mathisen, Karen Marie [Show all 10 contributors for this article] (2022). GRENSEVILT – reduksjon av grensebarrierer for skandinavisk viltforvaltning. Sveriges Lantbruksuniversitet. ISSN 9789157699398. Full text in Research Archive
• Wikenros, Camilla; Zimmermann, Barbara; Aronsson, Malin; Eriksen, Ane; Mathisen, Karen Marie & Persson, Jens [Show all 8 contributors for this article] (2022). GRENSEVILT – slutrapport till Interreg Sverige-Norge. Sveriges Lantbruksuniversitet. ISSN 9789157699381. Full text in Research Archive
• Aronsson, Malin; Persson, Jens; Zimmermann, Barbara; Märtz, Johanna; Wabakken, Petter & Heeres, Rick [Show all 7 contributors for this article] (2022). Järven i Inre Skandinaviens skogslandskap – områdesbruk, födoval och reproduktion. Sveriges Lantbruksuniversitet. ISSN 9789157699459. Full text in Research Archive
• Zimmermann, Barbara; Mathisen, Karen Marie; Ausilio, Giorgia; Sand, Håkan; Wikenros, Camilla & Eriksen, Ane [Show all 20 contributors for this article] (2022). Elgvandringer i grenseland med følger for skogbruk, jakt og rovdyr. Sveriges Lantbruksuniversitet. ISSN 9789157699411. Full text in Research Archive
• Sand, Håkan; Zimmermann, Barbara; Berg, Erik; Bramorska, Beata; Eriksen, Ane & Wikenros, Camilla [Show all 10 contributors for this article] (2022). Vandringsmönster hos GPS-försedda älgar i GRENSEVILT – konsekvenser för förvaltningen. Sveriges Lantbruksuniversitet. ISSN 9789157699404. Full text in Research Archive
• Eriksen, Ane; Willebrand, Maria Hörnell; Zimmermann, Barbara; Wikenros, Camilla; Åkesson, Mikael & Backer, Inge Lorange [Show all 16 contributors for this article] (2020). Assessment of the Norwegian part of the Scandinavian wolf population, phase 1. Høgskolen i Innlandet. Full text in Research Archive
• Tofastrud, Morten; Zimmermann, Barbara; Devineau, Olivier; Hessle, Anna; Aletengqimuke, S & Dickel, Lisa [Show all 10 contributors for this article] (2019). Beiteatferd, habitatseleksjon og produktivitet hos storfe på utmarksbeite. Universitetet i Innlandet. ISSN 9788283801576. Full text in Research Archive Show summary

  English: Political objectives of increased beef cattle production in Innlandet county will lead to an increased use of unproductive areas for cattle grazing, especially by specialized international beef cattle breeds. Cattle were bred to perform optimally on homogeneous, flat grasslands. In the forest, the habitat is highly heterogeneous, and grazing resources are patchily distributed and at lower concentrations. We have studied grazing behaviour, habitat selection and body growth of beef cattle in two study areas of southeast Norway. One area had a stocking density far below and the other well above the estimated grazing capacity. We used GPS-collars with activity sensors to monitor cattle movements every five or ten minutes throughout the summer grazing season. In addition, many cows and calves were weighted at release and homing. The cows used about one third of their time for grazing, but they adapted grazing time to the seasonally changing day length. In areas with a higher proportion of poor habitat, cattle had larger home ranges. Old summer farm meadows were the most preferred habitat type, followed by young forest stands younger than fifteen years. In the area of high stocking density, the animals additionally preferred habitat types of poorer grazing quality. For resting, the cattle preferred flat, grassy sites with canopy cover. Gain in body weight was highest for cows in the low stocking density area, while a higher proportion of cows at high stocking density lost weight during the grazing season. Earlymaturing breeds had highest increase in body weight in the area of low stocking density. Lactating cows had lower body weight gain, used smaller home ranges and spent more time grazing than dry cows. We conclude that international cattle breeds are suitable for beef production in the heterogenous environment of boreal forest pastures, given that the stocking density does not exceed the grazing capacity of the area. Further research should focus on positive and negative effects that grazing can have on biodiversity and other ecosystem services of the boreal forest, such as forestry, game harvest and tourism.

• Sand, Håkan; Mathisen, Karen Marie; Ausilio, Giorgia; Gicquel, M; Wikenros, Camilla & Månsson, Johan [Show all 10 contributors for this article] (2019). Kan forekomst av ulv redusere elgbeiteskader og øke tettheten av løvtrær? Utredning om ulv og elg del 4. Universitetet i Innlandet. ISSN 9788283800876. Full text in Research Archive Show summary

  Store rovdyr har nylig kommet tilbake til flere områder hvor de tidligere var utryddet, og dette har kastet nytt lys over deres potensielle effekter på biologisk mangfold og økosystemfunksjoner. Særlig effektene på lavere nivå i næringskjeden har fått mye oppmerksomhet. Store rovdyr kan påvirke vekst hos planter og biologisk mangfold, gjennom deres påvirkning på planteeteres tetthet og atferd (tri-trofiske kaskadeeffekter). Vi har undersøkt hypotesen om tri-trofiske kaskadeeffekter i det skandinaviske økosystemet, ved å fokusere på ulv, elg og planter som er viktige for elg, nemlig furu, og fire høyt selekterte løvtrær; rogn, osp, selje og eik (ROSE). Vi har belyst denne problemstillingen ved bruk av tre ulike datasett fra Norge og Sverige, hvor tetthet av elgmøkkhauger, beiteskader på furu, og tetthet av ROSE har blitt registrert over 1-13 år. Vi har fokusert på de seks fylkene i Sverige som har jevnlig forekomst av ulverevir, og områdene innenfor og like i nærheten av ulvesonen i Norge. Vi har analysert hvordan forekomst og tetthet av elg, beiteskader og tetthet av ROSE, varierer innenfor/utenfor ulverevir, og med antall år ulverevir har vært etablert i et område, i tillegg til en rekke andre miljøvariabler. Tetthet av elg og beiteskader viste enten ingen sammenheng med ulverevir, eller en svak økning innenfor ulverevir. Tetthet av ROSE viste oftest lavere tetthet innenfor ulverevir, men effekten var svak og varierte mellom datasett. Ulverevir forklarte generelt en veldig liten del av variasjonen i elgtetthet, beiteskader og ROSE. I stedet var hogstklasse, dekningsgrad av furu, bjørk og gran, avstand til eller tetthet av skogsbilveier, forekomst av gammelt elgbeite, høyde over havet og snødybde mye viktigere for å forklare variasjonen i elgtetthet, beite og ROSE. Vi fant heller ingen endring i habitatvalg hos elg, og dermed ingen støtte for en endring i atferd hos elg, avhengig av tilstedeværelse av ulv. Vi konkluderer med at våre resultater ikke gir støtte til hypotesen om at gjenkomst av ulv til Skandinavia fører til tri-trofiske kaskadeeffekter, med redusert beitetrykk på viktige plantearter. Våre resultat støtter ideen om at store rovdyr vil ha lavt potensiale for effekter på andre trofiske nivåer i områder med sterk menneskelig påvirkning, noe som er typisk for de områdene hvor store rovdyr nå kommer tilbake. Grunnen til dette er mest sannsynlig høyt jakttrykk på både rovdyr og byttedyr, og intensivt skogbruk og landbruk som gir grunnlag for høye byttedyrtettheter.

• Sand, Håkan; Wikenros, Camilla; Zimmermann, Barbara; Eriksen, Ane; Holen, Frode & Wabakken, Petter (2019). Predasjonsmønster hos enslige stasjonære ulver og ulver på vandring. Utredning om ulv og elg del 3. Høgskolen i Innlandet. Full text in Research Archive
• Liberg, Olof; Sand, Håkan; Wabakken, Petter; Wikenros, Camilla; Zimmermann, Barbara & Eriksen, Ane (2019). Beräkningar av beskattning av den Skandinaviska vargpopulationen 2020. Grimsö Wildlife Research Station, Department of Ecology, Swedish University of Agricultural Sciences. Full text in Research Archive
• Wikenros, Camilla; Zimmermann, Barbara; Sand, Håkan; Månsson, Johan; Maartmann, Erling & Eriksen, Ane [Show all 7 contributors for this article] (2019). Tildelt, sett og felt elg i forhold til ulveforekomst i Norge og Sverige. Utredning om ulv og elg del 1. Universitetet i Innlandet. ISSN 9788283800814. Full text in Research Archive Show summary

  Predasjon fra store rovdyr og jakt er de to viktigste faktorene som styrer dynamikken i mange hjorteviltpopulasjoner verden over. Gjenetableringen av ulv på den skandinaviske halvøya har møtt hard motstand fra mange hold, der ett argument har vært at ulveforekomst resulterer i en nedgang i elgstammen og reduserer det mulige jaktuttaket. Elgen er det viktigste byttedyret for ulven hele året, og ulvepredasjonen gjelder i hovedsak dyr som ellers kunne ha blitt felt under jakt. I denne rapporten presenterer vi hvordan ulven påvirker elgstammen og hvordan dette i sin tur påvirker avskytningen av elg i ulvens viktigste utbredelsesområde i Skandinavia ved å sammenligne områder uten og med ulik grad av ulveforekomst. Studien omfatter flere ulike datasett hvorav det mest omfattende spenner over en 32-års periode (fra 1986 i Norge og fra 1995 i Sverige til 2017). De mest detaljerte studiene av ulveforekomstens effekt på antall og sammensetning av kvoter, fellinger og observerte dyr ble gjennomført på data fra de seks siste årene. Vi har også sammenstilt hvilke jaktstrategier som er blitt utnyttet i den allmenne forvaltningen av elg og i forhold til forekomsten av ulv. Resultatet av denne studien viser at det har vært betydelig variasjon i avskytningen av elg i både Norge og Sverige de siste tiårene. En del av denne variasjonen kan forklares av den voksende ulvestammen. Utviklingen i elgstammen målt ved kvoter, fellinger og observerte dyr, samt effekten av ulv på disse variablene er likevel ulik mellom de to landene. Dette kan delvis forklares ved forskjeller i jaktstrategi, men også ved metodologiske ulikheter i innsamlingen av data. Denne studien viser at en endring i antall observerte dyr ett eller flere år ofte resulterte i en tilsvarende forandring i jaktuttaket neste år. Forekomst av ulv påvirket både antall og sammensetning av jaktuttak og observerte dyr. Det viste seg at ulven hadde en langtidseffekt på felte dyr per areal, mens påvirkningen på kjønns- og alderssammensetningen i sett elg var mer kortvarig. Data fra begge land viste at så vel kvoter som fellinger avtok den siste seksårs-perioden, og var koplet til ulveforekomst. Flertallet av analysene viste også at andre faktorer i kombinasjon med ulveforekomst har stor betydning for variasjonen i kvoter, felling og observerte dyr. Slike faktorer er breddegrad og andel kulturlandskap. Dette viser at endringer i både elgstammens størrelse, sammensetning og tilhørende jaktuttak styres av en kombinasjon av flere faktorer, som forvaltningsstrategi (jaktuttakets størrelse og sammensetning) og livsmiljø der predasjon fra store rovdyr er viktige elementer. Økt forekomst av ulv i Norge resulterte i at man kombinerte to jaktstrategier: 1. Man forsøkte å maksimere antallet felte dyr ved å øke andelen kalv i avskytningen. 2. Man reduserte jakttrykket på hunndyr i den hensikt å øke deres andel i populasjonen, altså skape en mer produktiv stamme. De samme strategier kunne observeres i Sverige, med den forskjell at andelen felte kalver de siste seks år var høyere enn i Norge, og uavhengig av om det fantes ulv. Dette viser at man i begge land innførte tiltak for å redusere den totale dødeligheten i elgstammen (redusert felling) så vel som for å maksimere produktiviteten (redusert felling av hunndyr). Den observerte andelen kalv i avskytningen tyder på at man uavhengig av ulveforekomst fant et kompromiss mellom å maksimere antall felte dyr og mengde kjøtt. Denne strategien var uavhengig (Sverige) eller ble styrket av den voksende ulvebestanden (Norge).

• Zimmermann, Barbara; Wikenros, Camilla; Sand, Håkan; Eriksen, Ane & Wabakken, Petter (2019). Elg i ulverevir: Predasjon og elgjakt. Utredning om ulv og elg del 2. Universitetet i Innlandet. ISSN 9788283800838. Full text in Research Archive Show summary

  I Skandinavia er ulvebestanden regulert av jakt. Dette medfører at ulvens påvirkning på byttedyrbestandene begrenses sammenlignet med uregulerte rovviltbestander og hovedsakelig konsentreres til områder med stasjonære, revirhevdende flokker og par. En måte å beregne effekten av ulvens predasjon på elgbestanden sammenlignet med jaktuttaket er å ta utgangspunkt i det enkelte ulverevir og beregne omfanget av ulvens predasjon ut fra kunnskap om ulvens predasjonstakt, størrelsen på reviret og elgtettheten. I den første delen av denne studien analyserer vi hvordan ulike elg- og ulverelaterte faktorer påvirker variasjonen i ulvens predasjonstrykk (andel av elgbestanden som tas av ulv) mellom ulike revir, samt hvordan dette henger sammen med eksisterende data for jaktuttak på revirnivå. I den andre delen av denne studien undersøker vi ulvenes tidsbruk innenfor sine revir og hvordan denne geografisk sammenfaller med fordelingen av 1) ulvedrepte byttedyr (elg) funnet i felt, samt 2) jaktuttaket og elgobservasjoner mellom de ulike jaktenhetene som inngår i reviret. Predasjonstrykket fra ulv på elgbestander i ulike ulverevir hadde ingen tydelig sammenheng med tetthet av elg, ulveflokkens størrelse, eller hvor ofte en ulveflokk tok elg (predasjonstakt). Derimot var predasjonstrykket sterkt avhengig av den kombinerte effekten av revirets størrelse og elgtetthet, som til sammen utgjør hvor mange elger som finnes i reviret. Omfanget av predasjonstrykket ble beregnet til gjennomsnittlig 7 – 8% av elgbestanden og varierte mellom 2 – 12% mellom revir. Motsvarende var jakttrykket beregnet til gjennomsnittlig 15 – 19%, med en variasjon på 8 – 33% mellom revir. Sett over hele året var jakttrykket gjennomsnittlig 2.4 – 3.5 ganger høyere enn predasjonstrykket, men jakttrykket var konsentrert til en kort periode i jakttiden, mens predasjonstrykket fordelte seg mer regelmessig over hele året. Jakttrykket hadde ikke noen sammenheng med antall elg i reviret eller predasjonstrykket, noe som tyder på at den kombinerte effekten av ulv og jakt ikke er kompensatorisk. En sammenligning av predasjonstrykket fra ulv og bjørn på elg i Alaska med dødeligheten fra jakt, ulv og bjørn i Skandinavia tyder på at en mindre andel av elgbestanden i Skandinavia dør årlig. Delstudie to viste 1) at ulvene fordeler tiden sin ulikt i de ulike delene av reviret, 2) at denne ujevne fordelingen blir spesielt sterk på sommeren i revir der ulvene har ynglet, 3) at fordelingen av ulvedrepte elger er ujevn og følger fordelingen av ulvenes områdebruk, men 4) at den ujevne fordelingen av ulvedrepte elger, i motsetning til ulvenes tidsbruk, ikke kan forklares med om ulvene har ynglet eller ikke, eller av årstid. En konsekvens av ulvenes ujevne områdebruk i reviret er at antallet felte elger per jaktareal var lavere i deler av reviret der ulvene tilbragte mer tid og der ulvene drepte flere elger. I gjennomsnittet var jaktuttaket 24% lavere i områder som ble mye brukt av ulv, sammenlignet med områder der ulvene tilbragte lite tid. Andelen felte kalver var hele 41% lavere i områder med høy sommeraktivitet sammenlignet med områder med lav sommeraktivitet, mens forskjellen i andelen felte kalver var på 33% mellom høy- og lavaktivitetsområder for hele året. Resultatene viser at det finnes en variasjon i jaktuttaket innenfor reviret som delvis kan forklares av ulvens områdebruk. Derimot fant vi ikke noen negativ sammenheng mellom sett elg per jegertime og ulvenes områdebruk eller fordeling av ulvedrepte elger. I noen tilfeller ble det til og med sett flere elger i områder med mye ulveaktivitet. Dette tyder på at det råder en kompleks sammenheng mellom ulvenes fordeling, predasjon, jaktuttak og sett elg i tid og rom på den geografiske skalaen av et ulverevir.

• Wabakken, Petter; Zimmermann, Barbara; Eriksen, Ane; Maartmann, Erling; Nordli, Kristoffer & Sanchez, David Carricondo [Show all 8 contributors for this article] (2018). Sesongkonflikter mellom mennesker og ulv i områder med snø og trekkelg: Ulv i Slettåsreviret, 2009-2018. Høgskolen i Innlandet. ISSN 9788283800760. Full text in Research Archive
• Nordli, Kristoffer; Zimmermann, Barbara; Wabakken, Petter; Eriksen, Ane; Sanchez, David Carricondo & Maartmann, Erling [Show all 8 contributors for this article] (2018). Ulvevalpers flokksamhold og områdebruk i Skandinavia. Utredning om ulv og bosetting del 2. Universitetet i Innlandet. ISSN 9788283800708. Full text in Research Archive Show summary

  English: The basic social unit in wolves consists of the territory-marking pair, and together with their offspring from the contemporary and/or previous litters they form a family group, commonly referred to as the wolf pack. The family group's grouping behavior and movement pattern depends on the wolves' year cycle. Observations in Scandinavia from long-term series of GPS data, and the National Monitoring Program indicates that the overall cohesion of the pack gradually dissolves during the winter. Using 15 years of data distributed over 17 pack years, 21 adult breeding pairs and 30 pups with simultaneous GPS positioning, we investigated wolf pack cohesion and pup dispersal from January to early July. We investigated the cohesion by using a regressive non-linear time series analysis (GAMM) with an added interaction term on time adjusted to appropriate reference levels. The cohesion was strongest in early winter but declining over time, characterized by wolf pups becoming increasingly independent from the rest of their family group, before most pups (75%) eventually disperse in the spring before 14 months of age. The distance between litter siblings and between pups and parents became significantly different from zero during the mating season and remained significant to after the birth of the subsequent litter. Overall the predicted variation in cohesion between siblings and between pups and their parents was large. In other words, pups could be separated from their parents and other siblings already in early winter, though not so often or over shorter time periods than later in winter-spring. In cases when the pups were separated from the parents, the pups’ habitat selection could be characterized as elusive, more concealed and further away from human infrastructure. Our results are a strong contrast to the common belief that members of a wolf pack allegedly operate as a strictly synchronized unit. This also may have implications for the annual wolf monitoring program in terms of calculating the number of wolves and reproductions, and for the planning and implementation of an ethical and effective license hunt on Scandinavian wolf family groups in the future.

• Eriksen, Ane; Zimmermann, Barbara; Wabakken, Petter; Sanchez, David Carricondo; Sand, Håkan & Wikenros, Camilla (2018). Spredningsulvers bevegelser i forhold til menneskelig infrastruktur. Utredning om ulv og bosetting del 3. Universitetet i Innlandet. ISSN 9788283800722. Full text in Research Archive Show summary

  English: We present the first telemetry study of the habitat use of dispersing wolves in Scandinavia. During dispersal, young wolves enter unfamiliar areas, and their travel routes often follow natural as well as anthropogenic landscape corridors such as valleys and roads. With average dispersal distances of 225 km (males) and 154 km (females), dispersing wolves can theoretically appear anywhere on the Scandinavian peninsula. During the period 2003-2017, 31 wolves were GPS collard inside their natal territories, and followed during part of or the entire period of natal dispersal. We used step selection functions and conditional logistic regressions to evaluate the movement of dispersing wolves relative to human settlement and infrastructure, during different seasons and times of day, and during displacement and at so-called clusters. We also looked for changes over time since the wolves left their natal territories. Based on the assumption that young dispersing wolves are inexperienced, relatively poor hunters, travel across large distances and into unfamiliar areas, we expected that these wolves would often get close to human settlements. We also expected that they would use roads for efficient transportation, but that they would adjust their activity in time and space to avoid encounters with humans. Finally, we expected that the proximity to human activity would decrease over time as the dispersing wolves got older, gained experience and became better hunters. Our results show that the dispersing wolves were generally evasive towards human infrastructure and avoided proximity to settlements, high building densities, and open areas. The dispersing wolves were particularly evasive when moving to places where they would spend longer periods of time at day beds or consuming food. During transportation the wolves selected higher densities of forest roads, seemingly regardless of season and time of day. During nocturnal transportation bouts they did not avoid main roads. Consistent with earlier studies we interpret this as the wolves using roads for efficient travel. The dispersing wolves showed stronger avoidance of roads, buildings and open areas during day than during night, probably to avoid humans. Their movement relative to human infrastructure was stable throughout the first nine months after leaving the natal territory. Our findings regarding the movement of dispersing wolves is consistent with existing knowledge about wolf behavior toward human infrastructure, and we did not find evidence that dispersing wolves as a group seek human activity more than resident wolves do. Nevertheless, we saw cases of individual dispersing wolves appearing in areas with very high levels of human activity, inside settlements and near large traffic arteries. Our interpretation of these results is that dispersing wolves generally avoid humans in the same way as resident wolves, and that e.g. roads are mainly used when the likelihood of encountering humans is low. Even so, wolves traveling across long distances will often have to cross major roads or densely populated areas where the risk of detection by humans is high.

• Zimmermann, Barbara; Wabakken, Petter; Eriksen, Ane; Maartmann, Erling; Sanchez, David Carricondo & Versluijs, Erik [Show all 8 contributors for this article] (2018). Slettåsulvenes atferd i forhold til menneskelig bosetting gjennom et helt år. Utredning om ulv og bosetting del 6. Universitetet i Innlandet. ISSN 9788283800784. Full text in Research Archive Show summary

  Etter at Slettåsulvene i 2009 etablerte revir i et område som har vært tomt for ynglende ulv i trolig over hundre år, har det vært en vedvarende konflikt om ulvers atferd i forhold til bosetting. Slettåsulvene ble av lokalt bosatte oppfattet som nærgående, og det var grunnen til at forvaltningen merket lederparet og fire valper med GPS-halsbånd i januar-februar 2017. Det Skandinaviske Ulveforskningsprosjektet SKANDULV fikk i oppdrag å analysere de regelmessige GPS-posisjonene tatt hver time eller hver fjerde time, studere ulvenes atferd ved å oppsøke posisjoner i felt og kartlegge fordelingen av elg og rådyr om vinteren og sommeren ved hjelp av systematiske møkktellinger. Vi har søkt etter kadaverrester og ulvespor på 45 % av alle 4-timersposisjoner til de voksne ulvene tatt i perioden 13.1.2017–10.1.2018. I området som Slettåsulvene brukte som revir gjennom 2017 (880 km2) er menneskelig bosetting konsentrert langs ved Osensjøen, i fire tettsteder og noen spredte enkelthus i de lavereliggende, tørre områdene, mens hytter finnes hovedsakelig enten lavt eller høyt i terrenget. Om vinteren var elgen konsentrert i atskilte områder, ofte nære hus, mens elgfordelingen var jevnere gjennom sommeren. Vi fant lite rådyrmøkk, nær hus om vinteren og langt fra hus om sommeren. Lederparet i Slettåsflokken oppholdt seg gjennomsnittlig nærmere hus enn forventet fra tilgjengelig habitat om vinteren, spesielt nattestid og i områder med høy elgtetthet, trolig for å jakte. Under yngling valgte ulvene stort sett hi- og valpeplasser langt fra bebyggelse, trolig for å unngå menneskelig forstyrrelse, men de fortsatte å jakte elg nærmere hus enn forventet, trolig fordi elgkyrne valgte kalvingshabitat i slike områder. Om høsten (oktober-november) valgte ulvene å jakte og oppholde seg langt fra bebodde hus, trolig for å unngå menneskelig forstyrrelse under elgjakt eller som en konsekvens av elgens habitatvalg. Når fjorårsvalpene gikk atskilt fra de voksne før utvandring fra oppvekstreviret, holdt de seg gjennomsnittlig lenger unna hus enn foreldreparet på samme tidspunkt, trolig fordi de ikke fulgte med på de voksnes jaktturer eller fordi de var mer varsomme overfor menneskelig forstyrrelse. Vi konkluderer med at sporobservasjoner av Slettåsulvene nær bebodde hus trolig skyldtes ulvenes jaktatferd i områder med god tilgang til elg og rådyr, og at ulvene passerte hus mens de jaktet eller var på vei mellom dagleie og kadaver heller enn at de oppholdt seg ved hus over lenger tid.

• Sanz-Perez, Ana; Milleret, Cyril Pierre; Fernandez, Andres Avelino Ordiz; Uzal, Antonio; Sanchez, David Carricondo & Eriksen, Ane [Show all 11 contributors for this article] (2018). Oppvekstrevirets effekt på habitatvalg hos voksne ulver. Utredning om ulv og bosetting del 4. Universitetet i Innlandet. ISSN 9788283800746. Full text in Research Archive Show summary

  English: We investigated whether the degree of natal exposure to anthropogenic influences could explain some of the variation in the selection of habitat with regard to anthropogenic factors in Scandinavian wolves. In the first part of the study we tested whether anthropogenic influences in the natal habitat might affect the choice of a breeding territory. Because dispersal distance may influence habitat similarity between natal and established territories, and mammals often show male-biased dispersal, we investigated the importance of dispersal distance and sex, and we tested alternative methods for defining habitat availability. In the second part of the study we evaluated whether the natal habitat may affect the habitat selection of wolves within their established territories, and we tested two competing hypotheses. A) Wolves exposed to a high degree of human activity in the natal territory may avoid human-related features within their established territory. B) Wolves exposed to a high degree of human activity in the natal territory may select for human related features in the established territory because of a certain degree of adaptation or habituation to human activity. We found that female, and to a smaller degree male wolves with short dispersal distances established in natal-like habitat types, regardless of the definition of habitat availability and habitat similarity. This was not the case for wolves that dispersed longer distances. We also found weak but consistent statistical evidence that wolf pairs in which the female had dispersed from a natal territory with a high degree of anthropogenic influence tended to select areas further away from humans within their established territory. The effects of the natal territory of the males was less clear. Part of the background for this evaluation was a concern among people living in wolf areas that wolves growing up in areas with high anthropogenic influence would seek human activity. On the contrary, our results suggest that wolves that have been exposed to larger degrees of anthropogenic influence in the natal territory may avoid proximity to human settlements and infrastructure inside their established territories. Our methodological approach may be useful for the continued wolf research and management in Scandinavia and elsewhere, both for the monitoring of whether wolves’ response to exposure to humans changes following changes in the wolf population or in the patterns of human settlement, but also to compare the response of wolves in Scandinavia to those in other parts of Europe, in which wolves are recolonizing areas of much higher anthropogenic influence.

• Liberg, Olof; Sand, Håkan; Wabakken, Petter; Wikenros, Camilla; Zimmermann, Barbara & Eriksen, Ane (2018). Beräkningar av beskattning av den Skandinaviska vargpopulationen. Grimsö Wildlife Research Station, Department of Ecology, Swedish University of Agricultural Sciences. Full text in Research Archive
• Sanchez, David Carricondo; Zimmermann, Barbara; Wabakken, Petter; Eriksen, Ane; Maartmann, Erling & Sanz-Perez, Ana [Show all 8 contributors for this article] (2018). Individuell atferd hos ulv overfor menneskelig infrastruktur i Skandinavia. Høgskolen i Innlandet. ISSN 9788283800678. Full text in Research Archive Show summary

  Ulv er kjent til å være en habitatgeneralist med ynglende bestander i mange ulike typer habitat fra arktiske strøk, boreale skoger, åpne landbruksområder og til tettbefolkede områder i varme strøk. En viktig faktor for ulvens habitatvalg er tilgang til byttedyr, og flere studier har vist at i områder hvor menneskelig infrastruktur overlapper med tilgang til byttedyr, utnytter ulven natten for å jakte. Ikke bare ytre faktorer kan påvirke ulvers habitatvalg. Ulveindivider har ulik personalitet formet av nedarvede og lærte egenskaper, og det kan derfor forventes en stor variasjon i hvordan ulike individer oppfører seg overfor menneskelig infrastruktur. Vi kategoriserte Skandinaviske ulvers individuell atferd langs en gradient fra unnvikende til oppsøkende, der unnvikende betyr unngåelse av infrastruktur og oppsøkende betyr preferanse for eller større bruk av infrastruktur enn forventet fra tilgjengeligheten. Vi brukte GPS-posisjoner fra 55 forsknings- og forvaltningsmerkede ulver i tidsperioden 2001-2017 i individuelle stegvalganalyser for å beskrive habitatvalget for hvert individ, for deretter å teste om variasjonen i habitatvalget kan forklares med årstid, tid på døgnet, alder, kjønn og andre individ- og revirspesifikke egenskaper. Ulvene viste oftest ikke noe spesiell seleksjonsmønster eller var unnvikende overfor infrastruktur, og oppsøkende atferd var sjeldent, med et unntak: mer enn halvparten av ulvene viste en oppsøkende atferd overfor skogsbilveier, trolig for bruk til energieffektiv forflytning. Ulvene foretrakk områder med veldig lav bygningstetthet. På dagtid var ulvene mer unnvikende, trolig for å unngå møter med folk, og på natten var de mindre unnvikende eller mer oppsøkende, trolig for å jakte på byttedyr. Ulvene, og spesielt tispene, var mer unnvikende på sommeren, trolig for å skjerme valpene på hi- og valpeplassene fra menneskelig forstyrrelse. Om vinteren var ulvene mindre unnvikende eller mer oppsøkende, trolig for å jakte elg og rådyr som oppholdt seg nær menneskelig infrastruktur når det er snødekke. Ulvene nord i norsk ulvesone og tilgrensende områder var mer oppsøkende enn ulvene lenger sør, trolig som en effekt av elgfordelingen i dalbunnene vinterstid i områder med elgtrekk. Vi fant ikke noe tydelig effekt av ulike individers eller flokkers særegenhet på deres atferdsmønster og antar derfor at ulvene i Skandinavia utviser en stor grad av fleksibilitet og tilpasningsevne i deres atferd overfor menneskelig infrastruktur.

• Veen, Bert van der; Mattisson, Jenny & Zimmermann, Barbara (2017). Where is the refrigerator? A spatial approach to wolverine (Gulo gulo) food hoarding behavior in Scandinavia. Universitetet i Innlandet. Full text in Research Archive Show summary

  Food hoarding helps animals to maintain their physical condition during times of food scarcity. Wolverines (Gulo gulo) inhabitat mostly environments of low productivity, and are known to hoard food for long-term periods, to store a single food item per food cache, and to spatially spread their food caches across their home range. Few studies have been dedicated to describing the wolverine’s food hoarding behavior. We identified 303 caches from 38 individuals in four study areas in Scandinavia, by exploring clusters of GPS positions during periods of four to eight weeks, in both summer and winter, with a total of 2090 monitoring days. During summer, it was easier to identify caches than in winter, when signs of caching activity are quickly covered up by snow. We studied the wolverine’s cache dispersion by means of a Complete Spatial Randomness test, identified their preferred caching habitat through resource selection modelling with conditional logistic regression, and applied linear mixed-effect models to study the caching distance between sources and caches. Wolverines hoarded food in both seasons and widely spaced their caches, occasionally clustering them in parts of their home range. Their optimal caching habitat was located in steep rugged terrain with plentiful vertical structure that offerered suitable micro-habitat. Wolverines did not cache further away from food sources to maximize cache habitat suitability. If a different carnivores species killed a prey animal, wolverines cached closer to a source. Ultimately the food hoarding behavior of wolverines is an adaptive response to survive periods of food scarcity.

• Wabakken, Petter; Maartmann, Erling; Eriksen, Ane; Zimmermann, Barbara; Flagstad, Øystein & Liberg, Olof [Show all 8 contributors for this article] (2017). Ulv som skadegjører på bufe, tamrein og hund i Norge: skadehistorikk og skadepotensiale i forhold til ulvens spredningsmønster. Universitetet i Innlandet. ISSN 9788283800036. Full text in Research Archive Show summary

  Since the reestablishment of the wolf population in Scandinavia during the 1980s, there have been conflicts related to the coexistence of wolves and people, mostly regarding loss of livestock and dogs, people’s fear, and competition for moose. To reduce conflicts, knowledge about wolf dispersal patterns is essential for management policies, but has thus far been studied very little. This is a first quantitative assessment of the history of, and potential for, livestock depredation by wolves in Norway. We have analyzed the dispersal patterns of Scandinavian wolves born in 1) fully Norwegian territories associated with today’s wolf zone, 2) trans-border territories, and 3) Swedish territories, respectively. Based on the established dispersal patterns, we analyzed wolf occurrence and potential for livestock depredation in Norway in general, as well as in the following regions: 1) Southern Norway outside of Hedmark County and the wolf zone, 2) Hedmark County, west of the Glomma River, and 3) Hedmark, east of Glomma and north of the wolf zone. Depredation on free-ranging domestic sheep is the major conflict concerning wolves and livestock in Norway. With the exception of two municipalities west of Oslo, there are no freeranging sheep in the wolf zone, and we conclude that young, solitary, dispersing wolves that immigrate to regions outside the wolf zone pose the greatest potential damage to free-ranging sheep in Norway. During the past 31 years, 2/3 of all wolf-killed sheep in Norway were in Hedmark County, and Rendalen Municipality in particular is suffering extensive and chronic losses, despite extensive culling of wolves. Most of the wolves immigrating to potential conflict areas in Southern Norway are born in Sweden. This is particularly true for those immigrating to Southern Norway outside of Hedmark and the wolf zone; among 56 wolves, 55 were Swedish-born, three were born in trans-border territories, and none were from the 41 wolf litters registered in fully Norwegian wolf territories associated with the wolf zone. Wolves from fully Norwegian territories east of Glomma almost always dispersed east into our neighboring countries. Only one out of 40 wolves which during a 20-year period were culled in Norway in direct association with attacks on livestock during the grazing season had dispersed from a fully Norwegian wolf territory associated with the wolf zone. We therefore conclude that the number of litters in fully Norwegian wolf territories seems to be of minimal importance for the extent of depredation on free-ranging sheep in Southern Norway as long as the reproductions happen in association with the wolf zone. Free-ranging cattle still graze both inside and outside the wolf zone, but the densities and the extent of wolf predation are low. During the period of 1995-2017, 113 dogs were classified as killed by wolves in Norway. The vast majority of these (107 dogs) were killed inside the wolf zone and/or inside Norwegian wolf territories. Resident, territorial wolves seem to be responsible for most of the attacks on dogs. Nevertheless, we found no relationship between the number of wolves or wolf territories in Norway and the number of dogs killed per year. Many territories had no attacks on dogs at all. Wolves in packs and pairs are an important factor in moose management, but the substantial decrease in moose populations in recent years in counties without resident wolves illustrates that factors other than wolves may be of major importance.

• Zimmermann, Barbara; Wabakken, Petter; Eriksen, Ane; Maartmann, Erling; Holen, Frode & Dahl, Espen R. [Show all 13 contributors for this article] (2017). Atferdsstudier på ulv i Slettås- og Osdalsreviret: Foreløpige resultater fra feltperioden januar–februar 2017. Høgskolen i Innlandet. ISSN 9788283800005. Full text in Research Archive Show summary

  Ulver i Slettåsreviret har i flere år blitt beskrevet som nærgående av lokalbefolkningen og media. Fem individer av flokken ble derfor merket med GPS-halsbånd i januar 2017, samtidig som ni ulver fra naboflokken i Osdalsreviret også ble GPS-merket. Det Skandinaviske Ulveforskningsprosjektet SKANDULV har deretter, med noen dagers forsinkelse for ikke å forstyrre, fulgt opp ulvene i felt for å kartlegge deres atferd og forflytningsmønster. Vi har oppsøkt de fleste plassene hvor ulvene har stoppet opp. Slike plasser karakteriseres av klynger med GPS-posisjoner, såkalte clustere, og i kombinasjon med sporsnø er det ofte mulig å se ulvenes atferd i slike områder. For de to ulveflokkene ble det etter fortløpende GIS-analyser undersøkt til sammen 5827 ulveposisjoner (83% av alle heltimesposisjonene) i felt, hvorav 5729 (98%) av posisjonene var fordelt klumpvis på totalt 362 clustere. Undersøkte GPS-posisjoner representerte en periode på 160 og 301 døgn for medlemmer av henholdsvis Slettåsflokken og Osdalsflokken, og GPS-posisjonene ble undersøkt av 7 peroner til sammen 67 dager i felt i perioden 7. februar – 2. mars 2017. På henholdsvis 8 (5%) og 14 (7%) clustere i Slettås- og Osdalsreviret fant vi rester etter elg som etter all sannsynlighet var drept av ulv, og på et cluster i Slettås var det rester etter det som kan ha vært en liten hund. Clustere med elgkadaver i Slettåsreviret lå gjennomsnittlig 1434 m (SE = 825 m) fra nærmeste bebodde hus. Dessuten ble matrester, ofte fra åteplasser, funnet på 12 clustere i Slettåsreviret. Denne typen cluster lå gjennomsnittlig nærmere bebodde hus enn andre typer av clustere. Det nærmeste matrester-cluster var på en åteplass besøkt av ulv 46 m fra hus. En foreløpig analyse av Slettåsulvenes GPS-posisjoner viser at ulvene foretrakk å bruke områder mellom 500 – 2000 m fra bebodde hus. Sett i forhold til andel areal med forskjellige avstander til bebodde hus brukte ulvene de nærmeste 500m fra hus mindre enn forventet på dagtid og omtrent som forventet om natten. Av de totalt 3095 posisjonene med 1-timesintervaller i Slettåsreviret var 21 (0,7%) nærmere enn 50 m fra bebodde hus, samtlige tatt om natten. Den nærmeste heltimesposisjonen påvist på dagtid var 98 m fra et hus. Den korte studieperioden begrenser muligheten til å trekke allmenne konklusjoner om ulvenes atferd, men vi kan se fra forflytningsmønsteret og clusterfunnene at mat trolig er en avgjørende faktor for ulvenes arealbruk. Fordelingen av elg i vinterkonsentrasjonsområder nær hus, men også tilgang til menneskeskapte åteplasser og dyp snø i høyden synes på denne årstiden å forklare mye av ulvenes forflytningsmønster. For en helhetlig konklusjon, økt allmenn kunnskap og framtidig adaptiv forvaltning av ulv er et anbefalt minimum å sammenligne Slettås- og Osdalsulvenes arealbruk med tilgjengelig tidligere års materiale fra ulver som er GPS-merket og observert i mer enn 50 andre Skandinaviske ulverevir.

• Panzacchi, Manuela; Marie, Stølhaug Christina & Zimmermann, Barbara (2016). Stress:does it matter? A review of mechanisms and fitness consequences of stress in large herbivores. Hedmark University College. Full text in Research Archive
• Chapron, Guillaume; Wikenros, Camilla; Liberg, Olof; Svensson, Linn; Åkesson, Mikael & Månsson, Johan [Show all 10 contributors for this article] (2015). Population estimates for the Scandinavian wolf population and sample-based monitoring. Viltskadecenter SLU. ISSN 9789186331795. Full text in Research Archive
• Linnell, John Durrus; Tveraa, Torkild; Swenson, Jon; Støen, Ole-Gunnar; Hansen, Inger & Andren, Henrik [Show all 21 contributors for this article] (2015). Kunnskapsstatus og kunnskapsbehov for forvaltning av rovvilt i Norge. Norsk institutt for naturforskning. ISSN 9788242628237. Full text in Research Archive Show summary

  Miljødirektoratet har bestilt denne rapporten for å identifisere kunnskapshull vedrørende rovvilt (gaupe, brunbjørn, ulv, jerv og kongeørn) i Skandinavia, med særlig fokus på situasjonen i Norge. Over 20 forskere fra ulike institusjoner i Norge og Sverige har bidratt til prosessen. Arbeidet har primært fokusert på naturvitenskapelige problemstillinger. De siste 30 årene har det vært gjennomført en omfattende forskning på store rovdyr i Skandinavia. Særlig har det vært gjennomført mange feltbaserte studier som har samlet inn demografiske data og atferdsdata gjennom bruk av telemetribaserte metoder. Parallelt har det vært en stor innsats på bestandsovervåking både ved bruk av tradisjonelle overvåkingsmetoder og genetiske analyser. I tillegg har det vært en betydelig forskningsinnsats på rovvilt innen andre disipliner. Forskningen har bidratt til et stort antall vitenskapelige publikasjoner – over 650 publikasjoner har kommet ut siden 1987, og det utgis nå rundt 50 publikasjoner i året. Den generelle biologien til disse artene er nå godt kjent og det finnes en god del kvantitative data for mange demografiske parametre, atferd og predator-byttedyrsammenhenger. Følgelig er de gjenstående kunnskapshullene fragmentert, spredt og varierer mellom artene. Kunnskap om demografi og områdebruk kommer fra et begrenset antall studieområder over begrensede tidsrom. Det er derfor til dels ukjent hvordan disse variablene varierer med endringer i bestandsstørrelse, klimatiske forhold og landskap. Et annet spørsmål er hvilke effekter den store avskytingen av rovdyr i Skandinavia har på atferdsmessige og demografiske parametre. Mesteparten av eksisterende kunnskap er knyttet til voksne individer. Vi har identifisert en rekke kunnskapshull knyttet til atferd og overlevelse hos juvenile – fra fødsel, gjennom spredningsfasen og fram til uavhengighet. Det er videre et behov for å forstå hvordan forskjellen i forvaltningsregime mellom Norge og Sverige påvirker våre felles rovviltbestander. Generelle trekk ved predasjonen på byttedyr er godt beskrevet. Det er imidlertid mange ubesvarte spørsmål knyttet til betydningen av romlig variasjon og byttedyrenes bevegelse (spredning og migrasjon) på predator-byttedyrdynamikken. Tilsvarende er det en del ubesvarte spørsmål knyttet til interaksjoner (f. eks. åtselspising / kleptoparasittisme) mellom store rovdyr, og hvordan disse påvirker byttedyrene. Alle feltstudiene har vært gjennomført i flerbrukslandskap som utnyttes både til skogbruk, jakt, friluftsliv og husdyrbeite. Den menneskelige aktiviteten kan ha stor innvirkning på store rovdyrs økologi og bør undersøkes. Eksempler på dette er effekten av skogsdrift på bjørn og betydningen av et stadig økende veinett på alle artene, inkludert interaksjonen med deres byttedyr. Innen temaene dyrehelse er det mange kunnskapshull å tette. Økt fokus på dyrevelferd, og klimaendringer som forventes å påvirke utbredelsen av parasitter og sykdommer, understreker behovet for å undersøke disse forholdene. Rovvilt, kunnskapsoversikt, kunnskapsbehov, rovvilt-beitedyr-konflikter, sau, tamrein, bjørn, gaupe, jerv, kongeørn, ulv, Large carnivores, Knowledge status, Knowledge needs, Carnivore - livestock conflict, Sheep, Semi-domestic reindeer, Brown bear, Eurasian Lynx, Golden Eagle, Wolverine, Wolf

• Zimmermann, Barbara; Wedul, Sari Jemiina; Johnsen, Kaja; Strømseth, Thomas Holm & Østerhus, Benedicte (2014). Hjort i Hedmark: Resultater fra GPS-merking 2002-2011. Høgskolen i Hedmark. ISSN 9788276719147. Full text in Research Archive Show summary

  Prosjektet Hjort i Hedmark var en del av samarbeidsprosjektet HjortAreal, som undersøkte hjortens arealbruk, økologisk bærekraft og hjort som næring i Møre & Romsdal og Sør-Trøndelag, Sogn i Hordaland, Buskerud, Haugalandet i Rogaland/Hordaland og Hedmark. Studieområdet i Hedmark hadde en unik posisjon i prosjektet fordi det representerte et ytterpunkt i forhold til klima, landskapsstruktur og tetthet av elg (Alces alces). Vi undersøkte Hedmarkshjortens trekkmønster og habitatbruk og da spesielt bruken av innmark og foringsplasser. Hjortemerking med GPS radiosendere begynte i Hedmark i 2002 og ble avsluttet vinteren 2011. Det ble totalt merket 51 hinder og vi fikk bra nok data fra 48 hinder. Sammenlignet med de andre studieområdene i HjortAreal-prosjektet var andelen trekkdyr høyest i Hedmark, der hele 96 % av hjortehindene var trekkdyr, og kun 4 % stasjonære. Også trekkavstandene var lengst i Hedmark. Hjorten trakk i gjennomsnitt 36.7 ± 7.5 km (2 SE) og tre av hjortene hadde sine respektive sommerområder rundt 100 km i luftlinje fra merkeplassen. Trekktidene er nokså sammenlignbare over hele Sør-Norge, med vårtrekket i løpet av april-mai og høsttrekket i august-september. Det var stor variasjon på størrelse av både vinter- og sommerområde. Gjennomsnittlig var vinterområde og sommerområde på henholdsvis 18.8 km2¬ (minst – størst 1.9–83.0 km2) og 22.9 km2 (1.7–98.0 km2). Avhengig av hvor hindene hadde sine vinterområder trakk de enten til høyereliggende eller lavereliggende sommerområde, eller vinter- og sommerområdene overlappet hverandre. Hjortens områdebruk ble styrt av menneskeskapte matressurser. Hele 10.5 % av posisjonene var knyttet til menneskeskapte konsentrasjoner av mat, enten de var innen 100 m av nærmeste foringsplass (3.1 %), eller på innmark (7.4 %). I månedsskifte februar-mars var en tredjedel av alle GPS-posisjonene knyttet til foringsplasser. Når bruken av foringsplasser avtok utover vinteren, fikk innmark mer og mer betydning. Innmarksarealet dekket gjennomsnittlig 3.4 ± 1.3 % (2 SE) av sommerleveområdene, og tilgangen varierte sterkt mellom hindene og områdene. Hindene oppholdt seg lenger enn forventet på innmark nattestid mens de brukte mindre tid enn forventet på innmark på dagtid. Samtlige foringsplasser i hoveddalføret langs Glomma i Stor-Elvdal og Rendalen kommuner hadde spor etter hjort, men ingen av foringsplassene hadde høye tettheter av både elg og hjort. Vi kan kun spekulere hvorvidt foring av elg har vært en medvirkende årsak for den nylige økningen i hjortebestanden i Hedmark. Vi har diskutert både plantefenologi-, konkurranse-, sosial barriere- og antipredator-hypotesen som en forklaring for Hedmarkhjortens trekkatferd med lange avstander og stor andel av trekkende dyr. Det kan tenkes et samspill av flere av disse hypotesene som grunnlag for at andel trekkdyr er høyest og trekkrutene er lengst i Hedmark sammenlignet med de andre studieområdene. Som delvis trekkende art kan hjorten utfordre forvaltningen. Fordi hjorten stort sett er tilbake på vinterområdene til jaktstart, kan kommune eller jaktvald være tilstrekkelig forvaltningsenhet, men ved en mer helthetlig forvaltning som tar hensyn til biologisk mangfold, vern av kalvingsområder og tiltak mot viltulykker, bør den romlige forvaltningsenheten være på regionalt nivå.

• Zimmermann, Barbara (2014). Predatory behaviour of wolves in Scandinavia. Universitetet i Innlandet. Full text in Research Archive Show summary

  The study of predatory behaviour is essential for understanding the interactions that may affect the behaviour and population dynamics of the species involved and consequent cascading ecosystem effects. In the case of large carnivores feeding on large ungulate prey, predation is strongly impacted by humans who alter the habitat of both predators and their prey, control population sizes, and compete with large carnivores for the same prey species. The objectives of this thesis were to 1) establish a robust method to estimate kill rates of wolves (Canis lupus); 2) estimate kill rates during both summer and winter; 3) relate kill rates to prey availability in winter (functional response) and to the energetic requirements of the wolf packs; and 4) to study the impact of roads on predatory and other behaviour of wolves. To meet these objectives, the Scandinavian Wolf Research Project (SKANDULV) fitted wolves with Global Positioning System (GPS) collars and searched for kill remains on clusters of GPS-positions during defined time periods in winter and summer. Winter prey densities of cervids were assessed by spring counts of faecal pellet groups. High variation in the spatio-temporal movement pattern of wolves in relation to kill sites made it difficult to predictably identify kill sites along the wolf trajectory without fieldwork. Consequently, field personnel checked all clusters, and in addition some single positions, to obtain robust estimates of kill rate. Human disturbance at kill sites may be one possible reason for the high variation in the handling of kills, with a relatively short handling time associated with kills located in areas with a high potential for human disturbance. During summer, wolves preferred to be at intermediate distances to roads while handling prey. This was probably the result of a trade-off between avoidance of human presence and the preference for roads while traveling. In most wolf territories, moose (Alces alces) was the pre-dominant prey species during both summer and winter. Kill rates, measured as the acquired edible biomass per pack, did not differ between summer (mean ± SE = 24.0 ± 3.0 kg/day/pack) and winter (28.2 ± 2.3 kg/day/pack), but wolves killed twice as many moose per unit time in summer (0.60 ± 0.05 moose/day/pack) as during winter (0.31 ± 0.04 moose/day/pack). This difference occurred because wolves selected for moose calves and in summer these provided significantly less biomass per moose kill than in winter. In both seasons, per capita kill rates were negatively associated with wolf pack size. This negative association, combined with a positive association between kill rate and prey availability, resulted in a predator-dependent functional response in winter. Small wolf packs (2 – 4 wolves) acquired more than three times as much biomass as required to cover the energetic needs of the pack, while large packs (7 – 9 wolves) in territories with low to intermediate prey abundance seemed to be food-limited during winter. The average annual kill rate was estimated at 120 moose per wolf pack (95% CI 100 – 144) in territories where moose accounted for the majority of the kills. Predation rates during winter, i.e. the proportion of the moose population killed by predators, were not related to wolf pack size or kill rates, but were negatively correlated with the abundance of moose or the moose-to-wolf ratio within the wolf territory. Wolves did not regulate, but rather limited, the highly managed moose population in Scandinavia, with wolf-induced mortality being mainly additive to other sources of mortality. If the wolf population is to be controlled as a measure to reduce predation on moose, the removal of non-breeding pairs and small family groups will likely have a stronger effect on moose kill rates than a reduction in the number of wolves in larger packs.

• Mysterud, Atle; Loe, Leif Egil; Meisingset, Erling; Zimmermann, Barbara; Hjeltnes, Arne William & Veiberg, Vebjørn [Show all 18 contributors for this article] (2011). Hjorten i det norske kulturlandskapet: arealbruk, bærekraft og næring. Universitetet i Oslo. ISSN 9788290934809. Full text in Research Archive
• Storaas, Torstein; Pedersen, Simen; Pedersen, Simen; Andreassen, Harry P.; Arnemo, Jon Martin & Dötterer, Michael [Show all 22 contributors for this article] (2008). Effekter av ulv på elgbestanden : da ulven kom og forsvant fra Koppangkjølen Effects of wolves on moose : when the wolf came and went from Koppangkjølen. Høgskolen i Hedmark. ISSN 9788276716474. Full text in Research Archive Show summary

  Fra 1997 til 2005 hevdet ulv revir på Koppangkjølen i Stor-Elvdal og Rendalen kommuner, Hedmark Fylke. Vi fulgte elgbestanden i området gjennom SETT ELG, takseringer fra helikopter og ved hjelp av radiomerkede elger. Vi har SETT ELG-data fra alle årene og gode data fra helikoptertakseringer og radiomerkede elger særlig fra 1999¿2001. Særlig mot slutten ble datainnsamlingen begrenset av prosjektøkonomien. Ulvenes antall og revirgrenser ble hver vinter kartlagt ved sporing på snø, og i 2004 ble tilleggsinformasjon innsamlet ved GPS-merking av både den revirhevdende tispa og hannen. Vi har beregnet det økonomiske tapet for jaktrettshaverne ved ulvens uttak av elg og analysert utviklingen av både reelle og potensielle inntekter fra småviltjakt og hytteutleie i ulvereviret. Basert på ulike deler av innsamlet datamateriale har vi produsert studentoppgaver på bachelor-, master- og ph.d.-nivå, i tillegg til diverse populærartikler og vitenskapelig publiserte arbeider. Her presenterer vi de viktigste resultatene fra de forskjellige arbeidene for å gi en samlet oversikt over hva vi lærte fra perioden med ulv på Koppangkjølen. During 1997 to 2005 a wolf territory was situated at Koppangkjølen in Stor-Elvdal and Rendalen municipalities, Hedmark County. We monitored the moose population in the area using a hunters¿ observation index called OBSERVED MOOSE, helicopter surveys, and by radiomarking moose. We have OBSERVED MOOSE data from all the years and good helicopter survey data and information from radiomarked moose, especially from 1999¿2001. Data collection was restricted in the later years by the project budget.The wolf pack numbers and territory boundaries were monitored each winter by snow tracking. In 2004 we got additional data by radiomarking both the female and male wolf territory holders. We estimated the landowner¿s economic loss by wolves killing moose and analyzed the development of the income from small game hunting and cabin renting within the wolf territory. Based on different parts of the data collected, our project has produced Bachelor-, Master-, and Ph.D.-theses, in addition to several popular articles and scientific papers. Herein we present the main results from our work and summarize the most important findings from the period with wolves at Koppangkjølen.

• Storaas, Torstein; Pedersen, Simen; Andreassen, Harry Peter; Arnemo, Jon Martin; Dötterer, Michael & Eriksen, Ane [Show all 21 contributors for this article] (2008). Effekter av ulv på elgbestanden: da ulven kom og forsvant fra Koppangkjølen. Høgskolen i Hedmark. ISSN 9788276716474. Full text in Research Archive Show summary

  English: During 1997 to 2005 a wolf territory was situated at Koppangkjølen in Stor-Elvdal and Rendalen municipalities, Hedmark County. We monitored the moose population in the area using a hunters’ observation index called OBSERVED MOOSE, helicopter surveys, and by radiomarking moose. We have OBSERVED MOOSE data from all the years and good helicopter survey data and information from radiomarked moose, especially from 1999–2001. Data collection was restricted in the later years by the project budget.The wolf pack numbers and territory boundaries were monitored each winter by snow tracking. In 2004 we got additional data by radiomarking both the female and male wolf territory holders. We estimated the landowner’s economic loss by wolves killing moose and analyzed the development of the income from small game hunting and cabin renting within the wolf territory. Based on different parts of the data collected, our project has produced Bachelor-, Master-, and Ph.D.-theses, in addition to several popular articles and scientific papers. Herein we present the main results from our work and summarize the most important findings from the period with wolves at Koppangkjølen. Within the Koppang wolf territory, a summer population of about 700 moose increased to more than 1 100 individuals due to migration in winter. The wolf pack size varied between 2 and 11 wolves. Of 135 radiomarked adult moose observed over 505 observed moose years, only 2 moose were classified as probably killed by wolves. However, many unmarked calves and yearlings were killed. It proved difficult to estimate exactly how many moose the wolves killed based on different indirect study methods on moose. The calf losses also increased with larger wolf packs and lower winter temperatures. The two years with good comparable data, we got inaccurate and varying estimates by the different methods with point estimates ranging from 62–175 moose per year. In our models we used a mean annual predation of 115 moose. This fits well with the yearly loss of 100–130 moose per wolf pack estimated by The Scandinavien wolf project (SKANDULV) by following GPS-marked wolves. Over 8 years, this mean level of predation would probably give a total loss of just below 1000 moose. Our models showed that the moose population would decrease dramatically with this level of predation if traffic accidents and harvest remained constant. Most likely, the predation on moose was even higher in the years when the wolf pack was at its largest and we had the best data. However, predation estimates based on OBSERVED MOOSE during these years were biased by a change in local harvest and hunting practices as a result of the wolf pack presence and the landowners’ interest in not reducing the moose population. Therefore, we based our estimate for these years on helicopter surveys, radiomarked moose and OBSERVED MOOSE from before the change in hunting practice, and estimated a mean predation loss of 160 moose a year within the wolf territory. If predation was this high in every year, the total loss would approach 1300 moose. When the old wolf female disappeared, the new territorial pair used a much larger area, and as a result the losses to wolves inside the old territory at Koppangkjølen were probably lower. A wolf pack hunts prey within its territory boundaries. However, as Koppangkjølen is a wintering area for migrating moose, the wolves could also kill moose that summered outside the territory when they moved into the territory during winter. We observed that the migrating cows lost a larger proportion of their calves in winter than cows resident year-round within the wolf territory. We also observed that the moose behaved slightly differently on supplementary feeding sites inside and outside the wolf territory. This indicates that local moose behavior may be affected by experience of wolf presence. The loss of moose to wolves is probably diluted across a larger area, neighboring the wolf territory because the wolves killed many calves of migrating moose. Furthermore net calf production was lower inside the territory and we suggest there was greater immigration than emigration, especially of young bulls. Our models indicated that the immigration of calf-loosing moose during winter reduced the losses to wolves of the resident moose population considerably. It proved difficult to estimate the loss of moose to wolves from hunter harvest data before and after the wolves established and disappeared. From the harvest data in the wolf territory 5 years before the wolves established, it seems as if the landowners reduced the harvest by 250 moose and that this reduction was sufficient to increase the moose population during the wolf period. However, the moose at Koppangkjølen is managed in 6 different management units with sometimes differing spatial and temporal goals. To determine the real losses during the wolf period, the harvest must be compared to the current goals of each management unit. In Koppang management unit (ca 1/6 of the wolf territory) the land owners reduced the harvest over 8 years by more than 300 moose to achieve their management goal of maintaining the population size. The other management units reduced their harvest by less. As the hunting practice and harvest changed inside the wolf territory, the hunter-based information from OBSERVED MOOSE also changed, most likely biasing the information. Hunter-based statistics on harvest and OBSERVED MOOSE are crude tools that have to be used with care to estimate losses to wolves. From our experience we conclude that moose losses to wolves will best be achieved by intensive studies of GPS-collared predators. As the local moose harvest was reduced, the calf losses to wolves were distributed over a larger area than the wolf territory because of wintering migration, and immigration was greater than emigration, the winter moose population within the wolf territory was not reduced despite many moose being killed by wolves. Thus the browsing pressure on pine plantations was not reduced. Because damage on forest regrowth may be substantial in high moose density areas, when wolves establish in an area we need models to predict when it is economically worth reducing the moose harvest and stabilizing the moose population, or on the contrary, when it will be best to reduce the moose population and invest in forest regeneration. If the mean annual wolf predation was as high as the years we had most wolves and best data, the wolf pack during the 8 year period reduced the income from moose hunting by 13 million NOK and gave a potential loss of 3 million from small game hunting and cabin renting. If so, total losses for the landowners within the wolf territory over 8 years may be close to 16 million NOK, with an annual loss of approximately 2 million NOK compared to the income if the wolves had settled in some other area. To reduce widespread illegal killing, and to achieve sustainable management and higher acceptance of wolves locally, a management strategy for the regional and national authorities could be to (after negotiations) also generously pay the local landowners and municipalities for economic losses other than loss of domestic animals where resident wolves are protected.

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