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.
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
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.
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.
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.
Understanding the distribution of biodiversity across the Earth is one of the most challenging questions in biology. Much research has been directed at explaining the species latitudinal pattern ...showing that communities are richer in tropical areas; however, despite decades of research, a general consensus has not yet emerged. In addition, global biodiversity patterns are being rapidly altered by human activities. Here, we aim to describe large‐scale patterns of species richness and diversity in terrestrial vertebrate scavenger (carrion‐consuming) assemblages, which provide key ecosystem functions and services. We used a worldwide dataset comprising 43 sites, where vertebrate scavenger assemblages were identified using 2,485 carcasses monitored between 1991 and 2018. First, we evaluated how scavenger richness (number of species) and diversity (Shannon diversity index) varied among seasons (cold vs. warm, wet vs. dry). Then, we studied the potential effects of human impact and a set of macroecological variables related to climatic conditions on the scavenger assemblages. Vertebrate scavenger richness ranged from species‐poor to species rich assemblages (4–30 species). Both scavenger richness and diversity also showed some seasonal variation. However, in general, climatic variables did not drive latitudinal patterns, as scavenger richness and diversity were not affected by temperature or rainfall. Rainfall seasonality slightly increased the number of species in the community, but its effect was weak. Instead, the human impact index included in our study was the main predictor of scavenger richness. Scavenger assemblages in highly human‐impacted areas sustained the smallest number of scavenger species, suggesting human activity may be overriding other macroecological processes in shaping scavenger communities. Our results highlight the effect of human impact at a global scale. As species‐rich assemblages tend to be more functional, we warn about possible reductions in ecosystem functions and the services provided by scavengers in human‐dominated landscapes in the Anthropocene.
Vertebrate scavenger (carrion‐consuming) species provide key ecosystem functions and services, but little is known about their large‐scale patterns of species richness and diversity. We found that scavenger assemblages in highly human‐impacted areas sustained the smallest number of scavenger species, suggesting human activity may be overriding other macroecological processes in shaping scavenger communities. As species‐rich assemblages tend to be more functional, we warn about possible reductions in ecosystem functions and the services provided by scavengers in human‐dominated landscapes in the Anthropocene.
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.
Trophic interactions are a fundamental topic in ecology, but we know little about how competition between apex predators affects predation, the mechanism driving top-down forcing in ecosystems. We ...used long-term datasets from Scandinavia (Europe) and Yellowstone National Park (North America) to evaluate how grey wolf (Canis lupus) kill rate was affected by a sympatric apex predator, the brown bear (Ursus arctos). We used kill interval (i.e. the number of days between consecutive ungulate kills) as a proxy of kill rate. Although brown bears can monopolize wolf kills, we found no support in either study system for the common assumption that they cause wolves to kill more often. On the contrary, our results showed the opposite effect. In Scandinavia, wolf packs sympatric with brown bears killed less often than allopatric packs during both spring (after bear den emergence) and summer. Similarly, the presence of bears at wolf-killed ungulates was associated with wolves killing less often during summer in Yellowstone. The consistency in results between the two systems suggests that brown bear presence actually reduces wolf kill rate. Our results suggest that the influence of predation on lower trophic levels may depend on the composition of predator communities.
Transboundary connectivity is a key component when conserving and managing animal species that require large areas to maintain viable population sizes. Wolves
Canis lupus
recolonized the Scandinavian ...Peninsula in the early 1980s. The population is geographically isolated and relies on immigration to not lose genetic diversity and to maintain long term viability. In this study we address (1) to what extent the genetic diversity among Scandinavian wolves has recovered during 30 years since its foundation in relation to the source populations in Finland and Russia, (2) if immigration has occurred from both Finland and Russia, two countries with very different wolf management and legislative obligations to ensure long term viability of wolves, and (3) if immigrants can be assumed to be unrelated. Using 26 microsatellite loci we found that although the genetic diversity increased among Scandinavian wolves (n = 143), it has not reached the same levels found in Finland (n = 25) or in Russia (n = 19). Low genetic differentiation between Finnish and Russian wolves, complicated our ability to determine the origin of immigrant wolves (n = 20) with respect to nationality. Nevertheless, based on differences in allelic richness and private allelic richness between the two countries, results supported the occurrence of immigration from both countries. A priori assumptions that immigrants are unrelated is non-advisable, since 5.8% of the pair-wise analyzed immigrants were closely related. To maintain long term viability of wolves in Northern Europe, this study highlights the potential and need for management actions that facilitate transboundary dispersal.
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