Directed endozoochorous dispersal by scavengers facilitate sexual reproduction in otherwise clonal plants at cadaver sites.

The regeneration niche of many plant species involves spatially and temporally unpredictable disturbances, called recruitment windows of opportunity. However, even species with clear dispersal adaptations such as fleshy berries may not successfully reach such elusive regeneration microsites. Ericaceous, berry-producing species in the northern hemisphere demonstrate this dispersal limitation. They are said to display a reproductive paradox owing to their lack of regeneration in apparently suitable microsites despite considerable investment in producing large quantities of berries.Cadavers generate vegetation-denuded and nutrient-rich disturbances termed cadaver decomposition islands (CDIs). Cadavers attract facultative scavengers with considerable capacity for endozoochorous seed dispersal. We hypothesize that CDIs facilitate recruitment in berry-producing ericaceous species due to endozoochorous dispersal directed toward favorable microsites with low competition.We examined seedling establishment within a permanent, semi-regular 10 × 10 m grid across an ungulate mass die-off on the Hardangervidda plateau in southeastern Norway. Competing models regarding the relative importance of factors governing recruitment were evaluated, specifically cadaver location (elevated seed rain) and microsite conditions (competition).We found that CDIs did facilitate seedling establishment, as cadaver density was the best predictor of seedling distribution. Other important factors governing seedling establishment such as percentage cover of soil and vascular plants alone were inadequate to explain seedling establishment. Synthesis: This study provides a novel understanding of sexual reproduction in species with cryptic generative reproduction. The directed nature of endozoochorous dispersal combined with long-distance dispersal abilities of medium to large vertebrate scavengers toward cadavers allows plants to exploit the advantageous but ephemeral resource provided by CDIs.

Harvest is associated with the disruption of social and fine-scale genetic structure among matrilines of a solitary large carnivore.

Harvest can disrupt wildlife populations by removing adults with naturally high survival. This can reshape sociospatial structure, genetic composition, fitness, and potentially affect evolution. Genetic tools can detect changes in local, fine-scale genetic structure (FGS) and assess the interplay between harvest-caused social and FGS in populations. We used data on 1614 brown bears, Ursus arctos, genotyped with 16 microsatellites, to investigate whether harvest intensity (mean low: 0.13 from 1990 to 2005, mean high: 0.28 from 2006 to 2011) caused changes in FGS among matrilines (8 matrilines; 109 females ≥4 years of age), sex-specific survival and putative dispersal distances, female spatial genetic autocorrelation, matriline persistence, and male mating patterns. Increased harvest decreased FGS of matrilines. Female dispersal distances decreased, and male reproductive success was redistributed more evenly. Adult males had lower survival during high harvest, suggesting that higher male turnover caused this redistribution and helped explain decreased structure among matrilines, despite shorter female dispersal distances. Adult female survival and survival probability of both mother and daughter were lower during high harvest, indicating that matriline persistence was also lower. Our findings indicate a crucial role of regulated harvest in shaping populations, decreasing differences among "groups," even for solitary-living species, and potentially altering the evolutionary trajectory of wild populations.

Causes and consequences of inverse density-dependent territorial behaviour and aggression in a monogamous mammal.

Territoriality is an important process shaping population dynamics, and the defence of a territory is crucial for individuals to increase the duration of territory occupancy and, consequently, reproductive success. However, little is known about how the frequency of territory intrusions and subsequent territorial behaviours and aggression by territory owners are affected by external factors, such as population density. This is important because it can affect mate change (the replacement of one pair member) and dispersal, a key ecological process. The aim of this study was to investigate the behavioural and spatial response of territory owners to intruder pressure as a function of population density in a territorial, monogamous mammal, the Eurasian beaver (Castor fiber). Using a combination of GPS technology, scent experiments, camera trap data and tail scar observations from an individual-based long-term study, we investigated the factors influencing spatial movement patterns by territory owners in response to a simulated intruder and the factors affecting territory intrusions. We found consistent inverse density-dependent patterns in territorial behaviours and evidence of conspecific aggression. At lower densities, territory owners detected more simulated intrusions, showed more territorial reactions and experienced increased conspecific aggression as indicated by tail scars, suggesting increased intruder pressure. Inverse density-dependent territorial behaviour and aggression suggest a potential mechanistic link between inverse density-dependent natal dispersal and mate change. At low population densities, increased dispersal amplifies intruder pressure, leading to the observed increases in territorial behaviours, conspecific aggression and previously observed mate turnover, which in turn might increase natal dispersal. Our study demonstrates how population density can affect the behaviour and space use of individuals, which is important for territory occupancy and fitness.

Heritability of head size in a hunted large carnivore, the brown bear (Ursus arctos).

Wild animal populations experience selection pressures from both natural and anthropogenic sources. The availability of extensive pedigrees is increasing along with our ability to quantify the heritability and evolvability of phenotypic traits and thus the speed and potential for evolutionary change in wild populations. The environment may also affect gene expressions in individuals, which may in turn affect the potential of phenotypic traits to respond to selection. Knowledge about the relationship between the genetic and environmental components of phenotypic variation is particularly relevant, given ongoing anthropogenically driven global change. Using a quantitative genetic mixed model, we disentangled the genetic and environmental components of phenotypic variance in a large carnivore, the brown bear (Ursus arctos). We combined a pedigree covering ~1,500 individual bears over seven generations with location data from 413 bears, as well as data on bear density, habitat characteristics, and climatic conditions. We found a narrow-sense heritability of 0.24 (95% CrI: 0.06-0.38) for brown bear head size, showing that the trait can respond to selection at a moderate speed. The environment contributed substantially to phenotypic variation, and we partitioned this into birth year (5.9%), nonadditive among-individual genetic (15.0%), and residual (50.4%) environmental effects. Brown bear head circumference showed an evolvability of 0.2%, which can generate large changes in the trait mean over some hundreds of generations. Our study is among the first to quantify heritability of a trait in a hunted large carnivore population. Such knowledge about the degree to which species experiencing hunting can respond to selection is crucial for conservation and to make informed management decisions. We show that including important environmental variables when analyzing heritability is key to understanding the dynamics of the evolutionary potential of phenotypic traits.

Sociodemographic factors modulate the spatial response of brown bears to vacancies created by hunting.

There is a growing recognition of the importance of indirect effects from hunting on wildlife populations, e.g. social and behavioural changes due to harvest, which occur after the initial offtake. Nonetheless, little is known about how the removal of members of a population influences the spatial configuration of the survivors. We studied how surviving brown bears (Ursus arctos) used former home ranges that had belonged to casualties of the annual bear hunting season in southcentral Sweden (2007-2015). We used resource selection functions to explore the effects of the casualty's and survivor's sex, age and their pairwise genetic relatedness, population density and hunting intensity on survivors' spatial responses to vacated home ranges. We tested the competitive release hypothesis, whereby survivors that increase their use of a killed bear's home range are presumed to have been released from intraspecific competition. We found strong support for this hypothesis, as survivors of the same sex as the casualty consistently increased their use of its vacant home range. Patterns were less pronounced or absent when the survivor and casualty were of opposite sex. Genetic relatedness between the survivor and the casualty emerged as the most important factor explaining increased use of vacated male home ranges by males, with a stronger response from survivors of lower relatedness. Relatedness was also important for females, but it did not influence use following removal; female survivors used home ranges of higher related female casualties more, both before and after death. Spatial responses by survivors were further influenced by bear age, population density and hunting intensity. We have shown that survivors exhibit a spatial response to vacated home ranges caused by hunting casualties, even in nonterritorial species such as the brown bear. This spatial reorganization can have unintended consequences for population dynamics and interfere with management goals. Altogether, our results underscore the need to better understand the short- and long-term indirect effects of hunting on animal social structure and their resulting distribution in space.

A "clearcut" case? Brown bear selection of coarse woody debris and carpenter ants on clearcuts.

Forest management alters habitat characteristics, resulting in various effects among and within species. It is crucial to understand how habitat alteration through forest management (e.g. clearcutting) affects animal populations, particularly with unknown future conditions (e.g. climate change). In Sweden, brown bears (Ursus arctos) forage on carpenter ants (Camponotus herculeanus) during summer, and may select for this food source within clearcuts. To assess carpenter ant occurrence and brown bear selection of carpenter ants, we sampled 6999 coarse woody debris (CWD) items within 1019 plots, of which 902 were within clearcuts (forests ⩽30 years of age) and 117 plots outside clearcuts (forests >30 years of age). We related various CWD and site characteristics to the presence or absence of carpenter ant galleries (nests) and bear foraging sign at three spatial scales: the CWD, plot, and clearcut scale. We tested whether both absolute and relative counts (the latter controlling for the number of CWD items) of galleries and bear sign in plots were higher inside or outside clearcuts. Absolute counts were higher inside than outside clearcuts for galleries (mean counts; inside: 1.8, outside: 0.8). CWD was also higher inside (mean: 6.8) than outside clearcuts (mean: 4.0). However, even after controlling for more CWD inside clearcuts, relative counts were higher inside than outside clearcuts for both galleries (mean counts; inside: 0.3, outside: 0.2) and bear sign (mean counts; inside: 0.03, outside: 0.01). Variables at the CWD scale best explained gallery and bear sign presence than variables at the plot or clearcut level, but bear selection was influenced by clearcut age. CWD circumference was important for both carpenter ant and bear sign presence. CWD hardness was most important for carpenter ant selection. However, the most important predictor for bear sign was the presence or absence of carpenter ant galleries. Bears had a high foraging "success" rate (⩾88%) in foraging CWD where galleries also occurred, which was assessed by summing CWD items with the concurrence of bear sign and galleries, divided by the sum of all CWD with bear sign. Clearcuts appeared to increase the occurrence of a relatively important summer food item, the carpenter ant, on Swedish managed forests for the brown bear. However, the potential benefit of this increase can only be determined from a better understanding of the seasonal and interannual variation of the availability and use of other important brown bear food items, berries (e.g. Vaccinium myrtillus and Empetrum spp.), as well as other primary needs for bears (e.g. secure habitat and denning habitat), within the landscape mosaic of managed forests.