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Abstract 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.

Abstract 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.

Abstract Estimating the contribution of demographic parameters to changes in population growth is essential for understanding why populations fluctuate. Integrated population models (IPMs) offer a possibility to estimate the contributions of additional demographic parameters, for which no data have been explicitly collected—typically immigration. Such parameters are often subsequently highlighted as important drivers of population growth. Yet, accuracy in estimating their temporal variation, and consequently their contribution to changes in population growth rate, has not been investigated. To quantify the magnitude and cause of potential biases when estimating the contribution of immigration using IPMs, we simulated data (using northern wheatear Oenanthe oenanthe population estimates) from controlled scenarios to examine potential biases and how they depend on IPM parameterization, formulation of priors, the level of temporal variation in immigration and sample size. We also used empirical data on populations with known rates of immigration: Soay sheep Ovis aries and Mauritius kestrel Falco punctatus with zero immigration and grey wolf Canis lupus in Scandinavia with near‐zero immigration. IPMs strongly overestimated the contribution of immigration to changes in population growth in scenarios when immigration was simulated with zero temporal variation (proportion of variance attributed to immigration = 63% for the more constrained formulation and real sample size) and in the wild populations, where the true number of immigrants was zero or near‐zero (kestrel 19.1%–98.2%, sheep 4.2%–36.1% and wolf 84.0%–99.2%). Although the estimation of the contribution of immigration in the simulation study became more accurate with increasing temporal variation and sample size, it was often not possible to distinguish between an accurate estimation from data with high temporal variation versus an overestimation from data with low temporal variation. Unrealistically, large sample sizes may be required to estimate the contribution of immigration well. To minimize the risk of overestimating the contribution of immigration (or any additional parameter) in IPMs, we recommend to: (a) look for evidence of variation in immigration before investigating its contribution to population growth, (b) simulate and model data for comparison to the real data and (c) use explicit data on immigration when possible.

Stunning by carbon dioxide (CO2) inhalation is controversial because it is associated with vigorous movements and behaviours which may or may not be conscious reactions. Furthermore, it is unknown whether some behaviours might indicate the transition into unconsciousness. Our study objective was to investigate the loss of consciousness during CO2 stunning by linking physiological variables (in particular pH, PaO2 and PaCO2) to the onset of observed behaviours. A total of 11 cross-bred pigs were studied. A tracheostomy tube, venous and arterial cannulae were placed under sevoflurane anaesthesia. After recovery from this, and a “wash out” period of at least 30 min, arterial blood samples were taken (and baseline values established) before 90–95% CO2 in medical air was administered through the tracheostomy tube. Subsequent behaviours were video-recorded and key physiological variables were evaluated using an anaesthetic monitor and the frequent sampling of arterial blood (albeit with inconsistent inter-sample intervals). After the study, behaviours were classified in an ethogram. At the onset of behaviours categorised as “vigorous movement extremities”, “opisthotonos” and “agonal gasping” pH values (range) were: 6.74–7.34; 6.66–6.96 and 6.65–6.87, while PaCO2 (kPa) was 4.6–42.2, 24.4–51.4 and 29.1–47.6. Based upon these values, we conclude that the pigs were probably unconscious at the onset of “opisthotonos” and “agonal gasping”, but some were probably conscious at the onset of “vigorous movements”.

Abstract Predation from large carnivores and human harvest are the two main mortality factors affecting 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 affected 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.

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.

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.

Knowledge about breeding biology is often incomplete in species with complex reproductive strategies. The brown bear Ursus arctos is a polygamous seasonal breeder inhabiting a wide variety of habitats and environmental conditions. We compiled information about brown bear breeding season dates from 36 study areas across their distribution range in the Palearctic and Nearctic regions and investigated how their breeding phenology relates to geographical factors (latitude, photoperiod, altitude and region). Brown bear matings were observed for 8 months, from April to November, with a peak in May–July. We found a 59-day difference in the onset of bear breeding season among study areas, with an average 2.3 days delay for each degree of latitude northwards. The onset of the breeding season showed a strong relationship with photoperiod and latitude, but not with region (i.e. Palearctic vs Nearctic) and altitude. First observations of bear mating occurred earlier in areas at lower latitudes. Photoperiod ranged between 14 and 18 hours at the beginning of the season for most of the study areas. The duration of the breeding season ranged from 25 to 138 days among study areas. None of the investigated factors was related to the length of the breeding season. Our results support the relevance of photoperiod to the onset of breeding, as found in other ursids, but not a shorter breeding season at higher latitudes, a pattern reported in other mammals. Our findings suggest a marked seasonality of bear reproductive behaviour, but also certain level of plasticity. Systematic field observations of breeding behaviour are needed to increase our knowledge on the factors determining mating behaviour in species with complex systems and how these species may adapt to climate change.

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 observation 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 reinforce that citizen-collected data are a useful wildlife monitoring tool that help understand ecological processes, including the effects of recovering carnivores on prey populations.

Laboratory mice are typically housed under extremely clean laboratory conditions, far removed from the natural lifestyle of a free-living mouse. There is a risk that this isolation from real-life conditions may lead to poor translatability and misinterpretation of results. We and others have shown that feral mice as well as laboratory mice exposed to naturalistic environments harbor a more diverse gut microbiota and display an activated immunological phenotype compared to hygienic laboratory mice. We here describe a naturalistic indoors housing system for mice, representing a farmyard-type habitat typical for house mice. Large open pens were installed with soil and domestic animal feces, creating a highly diverse microbial environment and providing space and complexity allowing for natural behavior. Laboratory C57BL/6 mice were co-housed in this system together with wild-caught feral mice, included as a source of murine microbionts. We found that mice feralized in this manner displayed a gut microbiota structure similar to their feral cohabitants, such as higher relative content of Firmicutes and enrichment of Proteobacteria. Furthermore, the immunophenotype of feralized mice approached that of feral mice, with elevated levels of memory T-cells and late-stage NK cells compared to laboratory-housed control mice, indicating antigenic experience and immune training. The dietary elements presented in the mouse pens could only moderately explain changes in microbial colonization, and none of the immunological changes. In conclusion, this system enables various types of studies using genetically controlled mice on the background of adaptation to a high diversity microbial environment and a lifestyle natural for the species.

This dataset article describes the data and sources used to model risks for the recolonizing wolf (Canis lupus) in Sweden and Norway in the article “Integrated spatially-explicit models predict pervasive risks to recolonizing wolves in Scandinavia from human-driven mortality” (Recio et al., 2018). Presences on wolf territories were used to model the potential distribution of the species. Presences of human-driven mortalities including traffic collisions, culling (protective/defensive, and licensed hunting), and illegal killing (i.e. poaching) were used to model predictions on the distribution of these mortalities. Sources for the independent variables used for the models are also described.

An amendment to this paper has been published and can be accessed via a link at the top of the paper.

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.

Circadian rhythms result from adaptations to biotic and abiotic environmental conditions that cycle through the day, such as light, temperature, or temporal overlap between interacting species. At high latitudes, close to or beyond the polar circles, uninterrupted midsummer daylight may pose a challenge to the circadian rhythms of otherwise nocturnal species, such as eagle owls Bubo bubo. By non‐invasive field methods, we studied eagle owl activity in light of their interactions with their main prey the water vole Arvicola amphibius, and their competitor the white‐tailed eagle Haliaeetus albicilla during continuous midsummer daylight on open, treeless islands in coastal northern Norway. We evaluated circadian rhythms, temporal overlap, exposure, and spatial distribution. The owls maintained a nocturnal activity pattern, possibly because slightly dimmer light around midnight offered favourable hunting conditions. The eagles were active throughout the 24‐h period as opposed to the strictly diurnal rhythm reported elsewhere, thus increasing temporal overlap and the potential for interference competition between the two avian predators. This may indicate an asymmetry, with the owls facing the highest cost of interference competition. The presence of eagles combined with constant daylight in this open landscape may make the owls vulnerable to interspecific aggression, and contrary to the available literature, eagle owls rarely exposed themselves visually during territorial calls, possibly to avoid detection by eagles. We found indications of spatial segregation between owls and eagles reflecting differences in main prey, possibly in combination with habitat‐mediated avoidance. Eagle owl vocal activity peaked in the evening before a nocturnal peak in visual observations, when owls were active hunting, consistent with the hypothesis of a dusk chorus in nocturnal bird species. The owls may have had to trade‐off between calling and foraging during the few hours around midnight when slightly dimmer light reduced the detection risk while also providing better hunting conditions

Brown bears (Ursus arctos) spend about half of the year in winter dens. In order to preserve energy, bears may select denning locations that minimize temperature loss and human disturbance. In expanding animal populations, demographic structure and individual behavior at the expansion front can differ from core areas. We conducted a non-invasive study of male brown bear den sites at the male-biased, low-density western expansion front of the Scandinavian brown bear population, comparing den locations to the available habitat. Compared to the higher-density population core in which intraspecific avoidance may affect den site selection of subordinate bears, we expected resource competition in the periphery to be low, and all bears to be able to select optimal den sites. In addition, bears in the periphery had access to free-ranging domestic sheep during summer. We found that males in the periphery denned on high-elevation slopes, probably providing good drainage, longer periods of consistent, insulating snow cover and fewer melting-freezing events. Forests were the principal denning habitat and no dens were found in alpine areas. The Scandinavian brown bears have a history of intense harvest, including culling at the den. This may have exerted a selection pressure to avoid denning in open alpine habitat which compared to forests provide little cover. The bears denned away from main roads and in steep, rugged terrain, probably limiting human access. The odds for finding a bear den decreased with increasing distance to the population core where females could be found. Previous studies have documented directed movement of male brown bears from the male-biased population periphery toward the core areas during the mating season. In this way, denning males may be trading off between low resource competition and access to sheep in the low-density periphery, and mating opportunities in the higher-density population core.

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

Research on large predator-prey interactions are often limited to the predators' primary prey, with the potential for prey switching in systems with multiple ungulate species rarely investigated. We evaluated wolf (Canis lupus) prey selection at two different spatial scales, i.e., inter- and intra-territorial, using data from 409 ungulate wolf-kills in an expanding wolf population in Scandinavia. This expansion includes a change from a one-prey into a two-prey system with variable densities of one large-sized ungulate; moose (Alces alces) and one small-sized ungulate; roe deer (Capreolus capreolus). Among wolf territories, the proportion of roe deer in wolf kills was related to both pack size and roe deer density, but not to moose density. Pairs of wolves killed a higher proportion of roe deer than did packs, and wolves switched to kill more roe deer as their density increased above a 1:1 ratio in relation to the availability of the two species. At the intra-territorial level, wolves again responded to changes in roe deer density in their prey selection whereas we found no effect of snow depth, time during winter, or other predator-related factors on the wolves' choice to kill moose or roe deer. Moose population density was only weakly related to intra-territorial prey selection. Our results show that the functional response of wolves on moose, the species hitherto considered as the main prey, was strongly dependent on the density of a smaller, alternative, ungulate prey. The impact of wolf predation on the prey species community is therefore likely to change with the composition of the multi-prey species community along with the geographical expansion of the wolf population.

Predators commonly select prey of inferior quality compared with the average animals found in the population. Consequently, predation may often be compensatory to other sources of mortality. We tested whether wolves (Canis lupus L., 1758) in Scandinavia selected moose in poor body condition by comparing mandibular marrow fat (MMF) of wolf-killed moose (Alces alces (L., 1758)) with harvested moose. Model selection analyses indicated that MMF levels were lower for wolf-killed moose compared with harvested moose, but an unbalanced sample design between years for the two causes of death (wolves and harvest) may have confounded a clear interpretation of the results. Nevertheless, corrected MMF levels (to 1 April) showed that a significantly (p = 0.006) higher proportion of the wolf-killed calves (18.5%) were below the level often assumed to reflect acute malnutrition ( Key words: compensatory predation, marrow fat, moose (Alces alces), prey selection, wolves (Canis lupus). Les predateurs selectionnent souvent des proies de qualite inferieure a celle des animaux moyens retrouves dans la population. En consequence, la predation peut s'averer etre compensatoire par rapport aux autres sources de mortalite. Nous verifions si les loups (Canis lupus L., 1758) en Scandinavie choisissent les elans en mauvaise condition corporelle en comparant la graisse de la moelle de la mandibule (MMF) chez des elans (Alces alces (L., 1758)) tues par les loups et d'autres elans recoltes. Des analyses de selection de modeles indiquent que les concentrations de MMF sont plus basses chez les elans tues par les loups que chez les autres elans recoltes, mais un plan d' echantillonnage non equilibre entre les annees pour les deux causes de mortalite (loups et recolte) peut avoir cause de la confusion dans l'interpretation des resultats. Neanmoins, les concentrations corrigees de MMF (jusqu'au 1 avril) montrent qu'une proportion significativement plus elevee (p = 0,006) de jeunes tues par les loups (18,5 %) se situent sous le niveau generalement presume refleter une malnutrition aigue ( Mots-cles : predation compensatoire, graisse de moelle, elan (Alces alces), selection des proies, loup (Canis lupus). [Traduit par la Redaction], Introduction The type of selection pattern by predators may be important for the dynamics of predator-prey populations for at least two reasons. First, selected prey individuals may con tribute differently [...]

Recent analyses have questioned the usefulness of heterozygosity estimates as measures of the inbreeding coefficient (f), a finding that may have dramatic consequences for the management of endangered populations. We confirm that f and heterozygosity is poorly correlated in a wild and highly inbred wolf population. Yet, our data show that for each level of f, it was the most heterozygous wolves that established themselves as breeders, a selection process that seems to have decelerated the loss of heterozygosity in the population despite a steady increase of f. The markers contributing to the positive relationship between heterozygosity and breeding success were found to be located on different chromosomes, but there was a substantial amount of linkage disequilibrium in the population, indicating that the markers are reflecting heterozygosity over relatively wide genomic regions. Following our results we recommend that management programs of endangered populations include estimates of both f and heterozygosity, as they may contribute with complementary information about population viability.

From November 1980 to May 1984, the Directorate for Wildlife and Freshwater Fish, Game Research Division has collected 285 reports of wolves Canis lupus in southeastern Norway. 64 reports have been verified as wolf, 69 have been rejected as non-wolf and 152 are unsolved. The main range for wolves in southeastern Norway was situated east of the river Glomma in Hedmark county. Wolves regularly crossed the border between Sweden and Norway, and the population is considered common to the two countries. Although it is difficult to draw any conclusions as to the total number of wolves within the study area, a minimum of three solitary adults was estimated in the first years of the study. Five wolves were the maximum number verified simultaneously in southeastern Norway.

Ref.: 90(3): 403-412. The affiliation for C. Wikenros should have been 'Grimso Wildlife Research Station, Department of Ecology, Swedish University of Agricultural Sciences, SE-730 91 Riddarhyttan, Sweden' and is correctly [...]