Publications by authors named "Håkan Sand"

33 Publications

Publisher Correction: Impact of a recolonizing, cross-border carnivore population on ungulate harvest in Scandinavia.

Sci Rep 2021 Apr 29;11(1):9602. Epub 2021 Apr 29.

Faculty of Applied Ecology, Agricultural Sciences and Biotechnology, Campus Evenstad, Inland Norway University of Applied Sciences, 2480, Koppang, Norway.

View Article and Find Full Text PDF

Download full-text PDF

Source
http://dx.doi.org/10.1038/s41598-021-89263-8DOI Listing
http://www.ncbi.nlm.nih.gov/pmc/articles/PMC8085227PMC
April 2021

Impact of a recolonizing, cross-border carnivore population on ungulate harvest in Scandinavia.

Sci Rep 2020 12 10;10(1):21670. Epub 2020 Dec 10.

Faculty of Applied Ecology, Agricultural Sciences and Biotechnology, Campus Evenstad, Inland Norway University of Applied Sciences, 2480, Koppang, Norway.

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.
View Article and Find Full Text PDF

Download full-text PDF

Source
http://dx.doi.org/10.1038/s41598-020-78585-8DOI Listing
http://www.ncbi.nlm.nih.gov/pmc/articles/PMC7730186PMC
December 2020

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

Sci Rep 2020 06 18;10(1):9941. Epub 2020 Jun 18.

Grimsӧ Wildlife Research Station, Department of Ecology, Swedish University of Agricultural Sciences, SE-730 91, Riddarhyttan, Sweden.

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

Download full-text PDF

Source
http://dx.doi.org/10.1038/s41598-020-66626-1DOI Listing
http://www.ncbi.nlm.nih.gov/pmc/articles/PMC7303184PMC
June 2020

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

Sci Rep 2019 04 25;9(1):6526. Epub 2019 Apr 25.

Faculty of Applied Ecology, Agricultural Sciences and Biotechnology, Inland Norway University of Applied Sciences, Evenstad, NO-2480, Koppang, Norway.

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.
View Article and Find Full Text PDF

Download full-text PDF

Source
http://dx.doi.org/10.1038/s41598-019-42835-1DOI Listing
http://www.ncbi.nlm.nih.gov/pmc/articles/PMC6484024PMC
April 2019

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

R Soc Open Sci 2018 Dec 12;5(12):181379. Epub 2018 Dec 12.

Faculty of Applied Ecology and Agricultural Sciences, Inland Norway University of Applied Sciences, Evenstad, 2480 Koppang, Norway.

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 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.
View Article and Find Full Text PDF

Download full-text PDF

Source
http://dx.doi.org/10.1098/rsos.181379DOI Listing
http://www.ncbi.nlm.nih.gov/pmc/articles/PMC6304128PMC
December 2018

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

Ecol Evol 2018 Dec 11;8(23):11450-11466. Epub 2018 Nov 11.

Grimsӧ Wildlife Research Station Department of Ecology Swedish University of Agricultural Sciences Riddarhyttan Sweden.

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

Download full-text PDF

Source
http://dx.doi.org/10.1002/ece3.4572DOI Listing
http://www.ncbi.nlm.nih.gov/pmc/articles/PMC6303696PMC
December 2018

Data to model risks for recolonizing wolves in Scandinavia through the integration of territory presence and human-driven mortalities.

Data Brief 2018 Oct 28;20:686-690. Epub 2018 Aug 28.

Grimsö Wildlife Research Station, Department of Ecology, Swedish University of Agricultural Sciences, SE-73091 Riddarhyttan, Sweden.

This dataset article describes the data and sources used to model risks for the recolonizing wolf () 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.
View Article and Find Full Text PDF

Download full-text PDF

Source
http://dx.doi.org/10.1016/j.dib.2018.08.060DOI Listing
http://www.ncbi.nlm.nih.gov/pmc/articles/PMC6129723PMC
October 2018

Genomic consequences of intensive inbreeding in an isolated wolf population.

Nat Ecol Evol 2018 Jan 20;2(1):124-131. Epub 2017 Nov 20.

Department of Evolutionary Biology, Evolutionary Biology Centre, Uppsala University, Norbyvägen 18D, SE-752 36, Uppsala, Sweden.

Inbreeding (mating between relatives) is a major concern for conservation as it decreases individual fitness and can increase the risk of population extinction. We used whole-genome resequencing of 97 grey wolves (Canis lupus) from the highly inbred Scandinavian wolf population to identify 'identical-by-descent' (IBD) chromosome segments as runs of homozygosity (ROH). This gave the high resolution required to precisely measure realized inbreeding as the IBD fraction of the genome in ROH (F ). We found a striking pattern of complete or near-complete homozygosity of entire chromosomes in many individuals. The majority of individual inbreeding was due to long IBD segments (>5 cM) originating from ancestors ≤10 generations ago, with 10 genomic regions showing very few ROH and forming candidate regions for containing loci contributing strongly to inbreeding depression. Inbreeding estimated with an extensive pedigree (F ) was strongly correlated with realized inbreeding measured with the entire genome (r  = 0.86). However, inbreeding measured with the whole genome was more strongly correlated with multi-locus heterozygosity estimated with as few as 500 single nucleotide polymorphisms, and with F estimated with as few as 10,000 single nucleotide polymorphisms, than with F . These results document in fine detail the genomic consequences of intensive inbreeding in a population of conservation concern.
View Article and Find Full Text PDF

Download full-text PDF

Source
http://dx.doi.org/10.1038/s41559-017-0375-4DOI Listing
January 2018

Fear or food - abundance of red fox in relation to occurrence of lynx and wolf.

Sci Rep 2017 08 22;7(1):9059. Epub 2017 Aug 22.

Forestry Research Institute of Sweden, Uppsala Science Park, SE-751 83, Uppsala, Sweden.

Apex predators may affect mesopredators through intraguild predation and/or supply of carrion from their prey, causing a trade-off between avoidance and attractiveness. We used wildlife triangle snow-tracking data to investigate the abundance of red fox (Vulpes vulpes) in relation to lynx (Lynx lynx) and wolf (Canis lupus) occurrence as well as land composition and vole (Microtus spp.) density. Data from the Swedish wolf-monitoring system and VHF/GPS-collared wolves were used to study the effect of wolf pack size and time since wolf territory establishment on fox abundance. Bottom-up processes were more influential than top-down effects as the proportion of arable land was the key indicator of fox abundance at the landscape level. At this spatial scale, there was no effect of wolf abundance on fox abundance, whereas lynx abundance had a positive effect. In contrast, at the wolf territory level there was a negative effect of wolves on fox abundance when including detailed information of pack size and time since territory establishment, whereas there was no effect of lynx abundance. This study shows that different apex predator species may affect mesopredator abundance in different ways and that the results may be dependent on the spatiotemporal scale and resolution of the data.
View Article and Find Full Text PDF

Download full-text PDF

Source
http://dx.doi.org/10.1038/s41598-017-08927-6DOI Listing
http://www.ncbi.nlm.nih.gov/pmc/articles/PMC5567382PMC
August 2017

Group or ungroup - moose behavioural response to recolonization of wolves.

Front Zool 2017 17;14:10. Epub 2017 Feb 17.

Department of Ecology, Grimsö Wildlife Research Station, Swedish University of Agricultural Sciences, SE-730 91 Riddarhyttan, Sweden.

Background: Predation risk is a primary motivator for prey to congregate in larger groups. A large group can be beneficial to detect predators, share predation risk among individuals and cause confusion for an attacking predator. However, forming large groups also has disadvantages like higher detection and attack rates of predators or interspecific competition. With the current recolonization of wolves () in Scandinavia, we studied whether moose () respond by changing grouping behaviour as an anti-predatory strategy and that this change should be related to the duration of wolf presence within the local moose population. In particular, as females with calves are most vulnerable to predation risk, they should be more likely to alter behaviour.

Methods: To study grouping behaviour, we used aerial observations of moose ( = 1335, where each observation included one or several moose) inside and outside wolf territories.

Results: Moose mostly stayed solitary or in small groups (82% of the observations consisted of less than three adult moose), and this behavior was independent of wolf presence. The results did not provide unequivocal support for our main hypothesis of an overall change in grouping behaviour in the moose population in response to wolf presence. Other variables such as moose density, snow depth and adult sex ratio of the group were overall more influential on grouping behaviour. However, the results showed a sex specific difference in social grouping in relation to wolf presence where males tended to form larger groups inside as compared to outside wolf territories. For male moose, population- and environmentally related variables were also important for the pattern of grouping.

Conclusions: The results did not give support for that wolf recolonization has resulted in an overall change in moose grouping behaviour. If indeed wolf-induced effects do exist, they may be difficult to discern because the effects from moose population and environmental factors may be stronger than any change in anti-predator behaviour. Our results thereby suggest that caution should be taken as to generalize about the effects of returning predators on the grouping behaviour of their prey.
View Article and Find Full Text PDF

Download full-text PDF

Source
http://dx.doi.org/10.1186/s12983-017-0195-zDOI Listing
http://www.ncbi.nlm.nih.gov/pmc/articles/PMC5316190PMC
February 2017

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

Proc Biol Sci 2017 02;284(1848)

Grimsö Wildlife Research Station, Department of Ecology, Swedish University of Agricultural Sciences, 730 91 Riddarhyttan, Sweden.

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

Download full-text PDF

Source
http://dx.doi.org/10.1098/rspb.2016.2368DOI Listing
http://www.ncbi.nlm.nih.gov/pmc/articles/PMC5310606PMC
February 2017

Mobility of moose-comparing the effects of wolf predation risk, reproductive status, and seasonality.

Ecol Evol 2016 Dec 21;6(24):8870-8880. Epub 2016 Nov 21.

Grimsö Wildlife Research Station Department of Ecology Swedish University of Agricultural Sciences Riddarhyttan Sweden.

In a predator-prey system, prey species may adapt to the presence of predators with behavioral changes such as increased vigilance, shifting habitats, or changes in their mobility. In North America, moose () have shown behavioral adaptations to presence of predators, but such antipredator behavioral responses have not yet been found in Scandinavian moose in response to the recolonization of wolves (). We studied travel speed and direction of movement of GPS-collared female moose ( = 26) in relation to spatiotemporal differences in wolf predation risk, reproductive status, and time of year. Travel speed was highest during the calving (May-July) and postcalving (August-October) seasons and was lower for females with calves than females without calves. Similarly, time of year and reproductive status affected the direction of movement, as more concentrated movement was observed for females with calves at heel, during the calving season. We did not find support for that wolf predation risk was an important factor affecting moose travel speed or direction of movement. Likely causal factors for the weak effect of wolf predation risk on mobility of moose include high moose-to-wolf ratio and intensive hunter harvest of the moose population during the past century.
View Article and Find Full Text PDF

Download full-text PDF

Source
http://dx.doi.org/10.1002/ece3.2598DOI Listing
http://www.ncbi.nlm.nih.gov/pmc/articles/PMC5192942PMC
December 2016

Prey Selection of Scandinavian Wolves: Single Large or Several Small?

PLoS One 2016 28;11(12):e0168062. Epub 2016 Dec 28.

Faculty of Applied Ecology and Agricultural Sciences, Hedmark University of Applied Sciences, Evenstad, Koppang, Norway.

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.
View Article and Find Full Text PDF

Download full-text PDF

Source
http://journals.plos.org/plosone/article?id=10.1371/journal.pone.0168062PLOS
http://www.ncbi.nlm.nih.gov/pmc/articles/PMC5193335PMC
July 2017

Let's stay together? Intrinsic and extrinsic factors involved in pair bond dissolution in a recolonizing wolf population.

J Anim Ecol 2017 Jan 28;86(1):43-54. Epub 2016 Sep 28.

Grimsö Wildlife Research Station, Department of Ecology, Swedish University of Agricultural Sciences, SE-730 91, Riddarhyttan, Sweden.

For socially monogamous species, breeder bond dissolution has important consequences for population dynamics, but the extent to which extrinsic or intrinsic population factors causes pair dissolution remain poorly understood, especially among carnivores. Using an extensive life-history data set, a survival analysis and competing risks framework, we examined the fate of 153 different wolf (Canis lupus) pairs in the recolonizing Scandinavian wolf population, during 14 winters of snow tracking and DNA monitoring. Wolf pair dissolution was generally linked to a mortality event and was strongly affected by extrinsic (i.e. anthropogenic) causes. No divorce was observed, and among the pair dissolution where causes have been identified, death of one or both wolves was always involved. Median time from pair formation to pair dissolution was three consecutive winters (i.e. approximately 2 years). Pair dissolution was mostly human-related, primarily caused by legal control actions (36·7%), verified poaching (9·2%) and traffic-related causes (2·1%). Intrinsic factors, such as disease and age, accounted for only 7·7% of pair dissolutions. The remaining 44·3% of dissolution events were from unknown causes, but we argue that a large portion could be explained by an additional source of human-caused mortality, cryptic poaching. Extrinsic population factors, such as variables describing the geographical location of the pair, had a stronger effect on risk of pair dissolution compared to anthropogenic landscape characteristics. Population intrinsic factors, such as the inbreeding coefficient of the male pair member, had a negative effect on pair bond duration. The mechanism behind this result remains unknown, but might be explained by lower survival of inbred males or more complex inbreeding effects mediated by behaviour. Our study provides quantitative estimates of breeder bond duration in a social carnivore and highlights the effect of extrinsic (i.e. anthropogenic) and intrinsic factors (i.e. inbreeding) involved in wolf pair bond duration. Unlike the effects of intrinsic and extrinsic factors that are commonly reported on individual survival or population growth, here we provide quantitative estimates of their potential effect on the social unit of the population, the wolf pair.
View Article and Find Full Text PDF

Download full-text PDF

Source
http://dx.doi.org/10.1111/1365-2656.12587DOI Listing
http://www.ncbi.nlm.nih.gov/pmc/articles/PMC5215671PMC
January 2017

Genetic rescue in a severely inbred wolf population.

Mol Ecol 2016 10 6;25(19):4745-56. Epub 2016 Sep 6.

Norwegian Institute for Nature Research, P.O. Box 5685 Sluppen, Trondheim, NO-7485, Norway.

Natural populations are becoming increasingly fragmented which is expected to affect their viability due to inbreeding depression, reduced genetic diversity and increased sensitivity to demographic and environmental stochasticity. In small and highly inbred populations, the introduction of only a few immigrants may increase vital rates significantly. However, very few studies have quantified the long-term success of immigrants and inbred individuals in natural populations. Following an episode of natural immigration to the isolated, severely inbred Scandinavian wolf (Canis lupus) population, we demonstrate significantly higher pairing and breeding success for offspring to immigrants compared to offspring from native, inbred pairs. We argue that inbreeding depression is the underlying mechanism for the profound difference in breeding success. Highly inbred wolves may have lower survival during natal dispersal as well as competitive disadvantage to find a partner. Our study is one of the first to quantify and compare the reproductive success of first-generation offspring from migrants vs. native, inbred individuals in a natural population. Indeed, our data demonstrate the profound impact single immigrants can have in small, inbred populations, and represent one of very few documented cases of genetic rescue in a population of large carnivores.
View Article and Find Full Text PDF

Download full-text PDF

Source
http://dx.doi.org/10.1111/mec.13797DOI Listing
http://www.ncbi.nlm.nih.gov/pmc/articles/PMC5054837PMC
October 2016

Sarcoptic mange in the Scandinavian wolf Canis lupus population.

BMC Vet Res 2016 Jul 27;12(1):156. Epub 2016 Jul 27.

Faculty of Applied Ecology and Agricultural Sciences, Hedmark University College, Campus Evenstad, N-2480, Koppang, Norway.

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 individual-based factors (sex, pack size) and the north-south gradient for seroprevalence suggests low probability of wolf-to-wolf transmission of S. scabiei in Scandinavia.
View Article and Find Full Text PDF

Download full-text PDF

Source
http://dx.doi.org/10.1186/s12917-016-0780-yDOI Listing
http://www.ncbi.nlm.nih.gov/pmc/articles/PMC4962404PMC
July 2016

Response of moose hunters to predation following wolf return in Sweden.

PLoS One 2015 8;10(4):e0119957. Epub 2015 Apr 8.

Grimsö Wildlife Research Station, Department of Ecology, Swedish University of Agricultural Sciences, Riddarhyttan, Sweden.

Background: Predation and hunter harvest constitute the main mortality factors affecting the size and dynamics of many exploited populations. The re-colonization by wolves (Canis lupus) of the Scandinavian Peninsula may therefore substantially reduce hunter harvest of moose (Alces alces), the main prey of wolves.

Methodology/principal Findings: We examined possible effects of wolf presence on hunter harvest in areas where we had data before and after wolf establishment (n = 25), and in additional areas that had been continuously exposed to wolf predation during at least ten years (n = 43). There was a general reduction in the total number of moose harvested (n = 31,827) during the ten year study period in all areas irrespective of presence of wolves or not. However, the reduction in hunter harvest was stronger within wolf territories compared to control areas without wolves. The reduction in harvest was larger in small (500-800 km2) compared to large (1,200-1,800 km2) wolf territories. In areas with newly established wolf territories moose management appeared to be adaptive with regard to both managers (hunting quotas) and to hunters (actual harvest). In these areas an instant reduction in moose harvest over-compensated the estimated number of moose killed annually by wolves and the composition of the hunted animals changed towards a lower proportion of adult females.

Conclusions/significance: We show that the re-colonization of wolves may result in an almost instant functional response by another large predator-humans-that reduced the potential for a direct numerical effect on the density of wolves' main prey, the moose. Because most of the worlds' habitat that will be available for future colonization by large predators are likely to be strongly influenced by humans, human behavioural responses may constitute a key trait that govern the impact of large predators on their prey.
View Article and Find Full Text PDF

Download full-text PDF

Source
http://journals.plos.org/plosone/article?id=10.1371/journal.pone.0119957PLOS
http://www.ncbi.nlm.nih.gov/pmc/articles/PMC4390345PMC
December 2015

Behavioral responses of wolves to roads: scale-dependent ambivalence.

Behav Ecol 2014 Nov 20;25(6):1353-1364. Epub 2014 Aug 20.

Department of Ecology, Grimsö Wildlife Research Station, Swedish University of Agricultural Science , SE-73091 Riddarhyttan , Sweden.

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 ( L.) from 14 territories in Scandinavia in relation to roads. We used resource and step selection functions, including >12000 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 km), they selected for roads when traveling, nearly doubling their travel speed. Breeding wolves moved the fastest. At the patch scale (10 km), house density rather than road density was a significant negative predictor of wolf patch selection. At the home range scale (approximately 1000 km), 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.
View Article and Find Full Text PDF

Download full-text PDF

Source
http://dx.doi.org/10.1093/beheco/aru134DOI Listing
http://www.ncbi.nlm.nih.gov/pmc/articles/PMC4235582PMC
November 2014

Predator-dependent functional response in wolves: from food limitation to surplus killing.

J Anim Ecol 2015 Jan 6;84(1):102-12. Epub 2014 Sep 6.

Faculty of Applied Ecology and Agricultural Sciences, Hedmark University College, Campus Evenstad, N-2480 Koppang, Norway.

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. 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. 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. 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. 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.
View Article and Find Full Text PDF

Download full-text PDF

Source
http://dx.doi.org/10.1111/1365-2656.12280DOI Listing
January 2015

Testing the risk of predation hypothesis: the influence of recolonizing wolves on habitat use by moose.

Oecologia 2014 Sep;176(1):69-80

Considered as absent throughout Scandinavia for >100 years, wolves (Canis lupus) have recently naturally recolonized south-central Sweden. This recolonization has provided an opportunity to study behavioral responses of moose (Alces alces) to wolves. We used satellite telemetry locations from collared moose and wolves to determine whether moose habitat use was affected by predation risk based on wolf use distributions. Moose habitat use was influenced by reproductive status and time of day and showed a different selection pattern between winter and summer, but there was weak evidence that moose habitat use depended on predation risk. The seemingly weak response may have several underlying explanations that are not mutually exclusive from the long term absence of non-human predation pressure: intensive harvest by humans during the last century is more important than wolf predation as an influence on moose behavior; moose have not adapted to recolonizing wolves; and responses may include other behavioral adaptations or occur at finer temporal and spatial levels than investigated.
View Article and Find Full Text PDF

Download full-text PDF

Source
http://dx.doi.org/10.1007/s00442-014-3004-9DOI Listing
September 2014

Decomposing risk: landscape structure and wolf behavior generate different predation patterns in two sympatric ungulates.

Ecol Appl 2013 Oct;23(7):1722-34

Norwegian Institute for Nature Research, P.O. Box 5685 Sluppen, NO-7485 Trondheim, Norway.

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 naive 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.
View Article and Find Full Text PDF

Download full-text PDF

Source
http://dx.doi.org/10.1890/12-1615.1DOI Listing
October 2013

Biomass flow and scavengers use of carcasses after re-colonization of an apex predator.

PLoS One 2013 23;8(10):e77373. Epub 2013 Oct 23.

Grimsö Wildlife Research Station, Department of Ecology, Swedish University of Agricultural Sciences, Riddarhyttan, Sweden.

Background: Reestablishment of apex predators influences the availability and distribution of biomass for scavengers and can therefore be an important agent for structuring species communities. We studied how the re-colonization of the Scandinavian Peninsula by wolves (Canis lupus) affected the amount and temporal variation in use of moose (Alces alces) carcasses.

Methodology/principal Findings: We compared the availability of biomass from remains at wolf kills with those killed by hunters, vehicle collisions and natural death. Movement-triggered cameras monitored patterns of use on wolf kills and remains from hunter harvest by scavengers (n = 15,276) in relation to time of year, available carcass biomass, time since the death of the moose and presence of wolves. Remains from hunter harvest were the largest food source for scavengers both within wolf territories (57%) and in areas without wolves (81%). The total annual biomass available were similar in areas with (25,648 kg) and without (24,289 kg) wolves. Presence of wolves lowered the peak biomass available from hunter harvest in October (20%) and increased biomass available during December to August (38-324% per month). The probability of scavengers being present decreased faster with time at remains from hunter harvest compared to wolf kills and both the probability of being present and the number of visits by scavengers to wolf kills increased as the amount of biomass available on the carcass increased.

Conclusions/significance: Wolves reduced the seasonal variation of biomass from moose carcasses and most important increased it during spring. Scavengers also visited wolf kills most frequently during spring when most scavenging species have young, which may lead to an increase in survival and/or reproductive success of scavengers within wolf territories. This applies both for abundant scavenging species that were the most frequent visitors at wolf kills and threatened scavengers with lower visit frequency.
View Article and Find Full Text PDF

Download full-text PDF

Source
http://journals.plos.org/plosone/article?id=10.1371/journal.pone.0077373PLOS
http://www.ncbi.nlm.nih.gov/pmc/articles/PMC3806759PMC
February 2015

Home range size variation in a recovering wolf population: evaluating the effect of environmental, demographic, and social factors.

Oecologia 2013 Nov 1;173(3):813-25. Epub 2013 May 1.

Norwegian Institute for Nature Research, Sluppen, P.O. Box 5685, Trondheim, 7485, Norway,

Home range size in mammals is a key ecological trait and an important parameter in conservation planning, and has been shown to be influenced by ecological, demographic and social factors in animal populations. Information on space requirements is especially important for carnivore species which range over very large areas and often come into direct conflict with human interest. We used long-term telemetry-location data from a recovering wolf population in Scandinavia to investigate variation in home range size in relation to environmental and social characteristics of the different packs. Wolves showed considerable variation in home range size, which ranged from 259 to 1,676 km(2). Although wolf density increased fourfold during the study period, we found no evidence that intraspecific competition influenced range size. Local variation in moose density, which was the main prey for most packs, did not influence wolf home range size. Home ranges increased with latitude and elevation and decreased with increased roe deer density. Although prey biomass alone did not influence range size, our data suggest that there is a correlation between habitat characteristics, choice of prey species and possible hunting success, which currently combine to shape home range size in Scandinavian wolves.
View Article and Find Full Text PDF

Download full-text PDF

Source
http://dx.doi.org/10.1007/s00442-013-2668-xDOI Listing
November 2013

Predicting the potential demographic impact of predators on their prey: a comparative analysis of two carnivore-ungulate systems in Scandinavia.

J Anim Ecol 2012 Mar 11;81(2):443-54. Epub 2011 Nov 11.

Norwegian Institute for Nature Research, PO Box 5685, Sluppen, NO-7485 Trondheim, Norway.

1. Understanding the role of predation in shaping the dynamics of animal communities is a fundamental issue in ecological research. Nevertheless, the complex nature of predator-prey interactions often prevents researchers from modelling them explicitly. 2. By using periodic Leslie-Usher matrices and a simulation approach together with parameters obtained from long-term field projects, we reconstructed the underlying mechanisms of predator-prey demographic interactions and compared the dynamics of the roe deer-red fox-Eurasian lynx-human harvest system with those of the moose-brown bear-gray wolf-human harvest system in the boreal forest ecosystem of the southern Scandinavian Peninsula. 3. The functional relationship of both roe deer and moose λ to changes in predation rates from the four predators was remarkably different. Lynx had the strongest impact among the four predators, whereas predation rates by wolves, red foxes, or brown bears generated minor variations in prey population λ. Elasticity values of lynx, wolf, fox and bear predation rates were -0·157, -0·056, -0·031 and -0·006, respectively, but varied with both predator and prey densities. 4. Differences in predation impact were only partially related to differences in kill or predation rates, but were rather a result of different distribution of predation events among prey age classes. Therefore, the age composition of killed individuals emerged as the main underlying factor determining the overall per capita impact of predation. 5. Our results confirm the complex nature of predator-prey interactions in large terrestrial mammals, by showing that different carnivores preying on the same prey species can exert a dramatically different demographic impact, even in the same ecological context, as a direct consequence of their predation patterns. Similar applications of this analytical framework in other geographical and ecological contexts are needed, but a more general evaluation of the subject is also required, aimed to assess, on a broader systematic and ecological range, what specific traits of a carnivore are most related to its potential impact on prey species.
View Article and Find Full Text PDF

Download full-text PDF

Source
http://dx.doi.org/10.1111/j.1365-2656.2011.01928.xDOI Listing
http://www.ncbi.nlm.nih.gov/pmc/articles/PMC3440569PMC
March 2012

Shoot, shovel and shut up: cryptic poaching slows restoration of a large carnivore in Europe.

Proc Biol Sci 2012 Mar 17;279(1730):910-5. Epub 2011 Aug 17.

Grimsö Wildlife Research Station, Department of Ecology, Swedish University of Agricultural Sciences, 73091 Riddarhyttan, Sweden.

Poaching is a widespread and well-appreciated problem for the conservation of many threatened species. Because poaching is illegal, there is strong incentive for poachers to conceal their activities, and consequently, little data on the effects of poaching on population dynamics are available. Quantifying poaching mortality should be a required knowledge when developing conservation plans for endangered species but is hampered by methodological challenges. We show that rigorous estimates of the effects of poaching relative to other sources of mortality can be obtained with a hierarchical state-space model combined with multiple sources of data. Using the Scandinavian wolf (Canis lupus) population as an illustrative example, we show that poaching accounted for approximately half of total mortality and more than two-thirds of total poaching remained undetected by conventional methods, a source of mortality we term as 'cryptic poaching'. Our simulations suggest that without poaching during the past decade, the population would have been almost four times as large in 2009. Such a severe impact of poaching on population recovery may be widespread among large carnivores. We believe that conservation strategies for large carnivores considering only observed data may not be adequate and should be revised by including and quantifying cryptic poaching.
View Article and Find Full Text PDF

Download full-text PDF

Source
http://dx.doi.org/10.1098/rspb.2011.1275DOI Listing
http://www.ncbi.nlm.nih.gov/pmc/articles/PMC3259920PMC
March 2012

Building a mechanistic understanding of predation with GPS-based movement data.

Philos Trans R Soc Lond B Biol Sci 2010 Jul;365(1550):2279-88

Department of Biological Sciences, University of Alberta, , Edmonton, Alberta, Canada.

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.
View Article and Find Full Text PDF

Download full-text PDF

Source
http://dx.doi.org/10.1098/rstb.2010.0077DOI Listing
http://www.ncbi.nlm.nih.gov/pmc/articles/PMC2894956PMC
July 2010

Summer kill rates and predation pattern in a wolf-moose system: can we rely on winter estimates?

Oecologia 2008 May 13;156(1):53-64. Epub 2008 Feb 13.

Grimsö Wildlife Research Station, Department of Ecology, Swedish University of Agricultural Sciences, 73091 Riddarhyttan, Sweden.

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.
View Article and Find Full Text PDF

Download full-text PDF

Source
http://dx.doi.org/10.1007/s00442-008-0969-2DOI Listing
May 2008

Selection for heterozygosity gives hope to a wild population of inbred wolves.

PLoS One 2006 Dec 20;1:e72. Epub 2006 Dec 20.

Department of Animal Ecology, Lund University, Lund, Sweden.

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.
View Article and Find Full Text PDF

Download full-text PDF

Source
http://journals.plos.org/plosone/article?id=10.1371/journal.pone.0000072PLOS
http://www.ncbi.nlm.nih.gov/pmc/articles/PMC1762340PMC
December 2006

Severe inbreeding depression in a wild wolf (Canis lupus) population.

Biol Lett 2005 Mar;1(1):17-20

Grimsö Wildlife Research Station, Department of Conservation Biology, Swedish University of Agricultural Science, SE-73091 Riddarhyttan, Sweden.

The difficulty of obtaining pedigrees for wild populations has hampered the possibility of demonstrating inbreeding depression in nature. In a small, naturally restored, wild population of grey wolves in Scandinavia, founded in 1983, we constructed a pedigree for 24 of the 28 breeding pairs established in the period 1983-2002. Ancestry for the breeding animals was determined through a combination of field data (snow tracking and radio telemetry) and DNA microsatellite analysis. The population was founded by only three individuals. The inbreeding coefficient F varied between 0.00 and 0.41 for wolves born during the study period. The number of surviving pups per litter during their first winter after birth was strongly correlated with inbreeding coefficients of pups (R2=0.39, p<0.001). This inbreeding depression was recalculated to match standard estimates of lethal equivalents (2B), corresponding to 6.04 (2.58-9.48, 95% CI) litter-size-reducing equivalents in this wolf population.
View Article and Find Full Text PDF

Download full-text PDF

Source
http://dx.doi.org/10.1098/rsbl.2004.0266DOI Listing
http://www.ncbi.nlm.nih.gov/pmc/articles/PMC1629062PMC
March 2005

Cross-continental differences in patterns of predation: will naive moose in Scandinavia ever learn?

Proc Biol Sci 2006 Jun;273(1592):1421-7

Grimsö Research Station, Swedish University of Agricultural Sciences, 730 91 Riddarhyttan, Sweden.

Predation has been recognized as a major selective force in the evolution of behavioural characteristics of mammals. As a consequence of local predator extinction, prey may lose knowledge about natural predators but usually express behavioural adjustments after return of predators. Human harvest may replace natural predation but prey selection may differ from that of natural predators leading to a change in the behavioural response of prey. We show that hunting success (HS) of re-colonizing wolves (Canis lupus) on moose (Alces alces) in Scandinavia was higher than reported in North America, where moose have been continuously exposed to wolves and grizzly bears. We found no evidence that moose expressed behavioural adjustments that lowered the HS of wolves in territories that had been occupied by wolves for up to 21 years. Moose behaviour towards wolves and humans typically differs in Scandinavia compared to North America. We explain the differences found to be caused by variation in predation pressure by large carnivores and the rate, and mode, of human harvest during the twentieth century.
View Article and Find Full Text PDF

Download full-text PDF

Source
http://dx.doi.org/10.1098/rspb.2005.3447DOI Listing
http://www.ncbi.nlm.nih.gov/pmc/articles/PMC1560300PMC
June 2006