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"Frank, Shane C"
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Ontogeny shapes individual dietary specialization in female European brown bears (Ursus arctos)
Individual dietary specialization, where individuals occupy a subset of a population’s wider dietary niche, is a key factor determining a species resilience against environmental change. However, the ontogeny of individual specialization, as well as associated underlying social learning, genetic, and environmental drivers, remain poorly understood. Using a multigenerational dataset of female European brown bears (Ursus arctos) followed since birth, we discerned the relative contributions of environmental similarity, genetic heritability, maternal effects, and offspring social learning from the mother to individual specialization. Individual specialization accounted for 43% of phenotypic variation and spanned half a trophic position, with individual diets ranging from omnivorous to carnivorous. The main determinants of dietary specialization were social learning during rearing (13%), environmental similarity (5%), maternal effects (11%), and permanent between-individual effects (9%), whereas the contribution of genetic heritability (3%) was negligible. The trophic position of offspring closely resembled the trophic position of their mothers during the first 3–4 years of independence, but waned with increasing time since separation. Our study shows that social learning and maternal effects were more important for individual dietary specialization than environmental composition. We propose a tighter integration of social effects into studies of range expansion and habitat selection under global change.
Journal Article
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.
Journal Article
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.
Journal Article
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.
Journal Article
Secondary forest floristic composition, structure, and spatial pattern in subtropical China
Secondary evergreen broadleaved forests are precious remnants for biodiversity conservation and templates for sustainable management of natural forests in subtropical China. Floristic composition, size structure, and spatial pattern of dominant tree species have been investigated for a subtropical secondary evergreen broadleaved forest in the Huitong Yingzuijie National Forest Reserve, Hunan, China. The location of all trees greater than 4 cm in diameter at breast height (DBH) were mapped within a 0.96-ha plot in which species, DBH, and total tree height were recorded. Ripley's K(t) function was used to analyze spatial patterns and associations. The secondary forest consisted of 74 tree species and 1,596 stems per hectare. A reverse-J shaped DBH classes distribution was observed for all stems and trees of later seral species whereas trees of earlier successional species were distributed irregularly. Significant aggregated spatial patterns were observed for all trees within the forest and for conspecific trees of each dominant species. This result, and a repulsive spatial pattern for interspecific trees of Choerospondias axillaries and Cyclobalanopsis glauca against other dominant tree species, support segregation hypothesis. Contributions of seed dispersal, topographic heterogeneity, and competition to spatial patterns of conspecific trees vary depending on tree species. Attractive spatial pattern among interspecific trees of Liquidambar fortunei, Liquidambar formosana, and Pinus massoniana reflects stochastic colonization of pioneer tree species and a facilitation relationship. Although deciduous species are long-lived and persist over long successional processes, they will eventually be replaced by late seral evergreen species within the secondary forest if no disturbance events occur.
Journal Article
Sociodemographic factors modulate the spatial response of brown bears to vacancies created by hunting
1. 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. 2. 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. 3. 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. 4. 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. 5. 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.
Journal Article
Indirect effects of bear hunting: a review from Scandinavia
Harvest by means of hunting is a commonly used tool in large carnivore management. To evaluate the effects of harvest on populations, managers usually focus on numerical or immediate direct demographic effects of harvest mortality on a population's size and growth. However, we suggest that managers should also give consideration to indirect and potential evolutionary effects of hunting (e.g., the consequences of a change in the age, sex, and social structure), and their effects on population growth rate. We define “indirect effects” as hunting-induced changes in a population, including human-induced selection, that result in an additive change to the population growth rate “lambda” beyond that due to the initial offtake from direct mortality. We considered 4 major sources of possible indirect effects from hunting of bears: (1) changes to a population's age and sex structure, (2) changes to a population's social structure, (3) changes in individual behavior, and (4) human-induced selection. We identified empirically supported, as well as expected, indirect effects of hunting based primarily on >30 years of research on the Scandinavian brown bear (Ursus arctos) population. We stress that some indirect effects have been documented (e.g., habitat use and daily activity patterns of bears change when hunting seasons start, and changes in male social structure induce sexually selected infanticide and reduce population growth). Other effects may be more difficult to document and quantify in wild bear populations (e.g., how a younger age structure in males may lead to decreased offspring survival). We suggest that managers of bear and other large carnivore populations adopt a precautionary approach and assume that indirect effects do exist, have a potential impact on population structure, and, ultimately, may have an effect on population growth that differs from that predicted by harvest models based on direct effects alone.
Journal Article
Water body type and group size affect the flight initiation distance of European waterbirds
This is an open access article distributed under the terms of the Creative Commons Attribution License, which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Journal Article
Hunting promotes spatial reorganization and sexually selected infanticide
Harvest can affect the ecology and evolution of wild species. The removal of key individuals, such as matriarchs or dominant males, can disrupt social structure and exacerbate the impact of hunting on population growth. We do not know, however, how and when the spatiotemporal reorganization takes place after removal and if such changes can be the mechanism that explain a decrease in population growth. Detailed behavioral information from individually monitored brown bears, in a population where hunting increases sexually selected infanticide, revealed that adult males increased their use of home ranges of hunter-killed neighbors in the second year after their death. Use of a hunter-killed male’s home range was influenced by the survivor’s as well as the hunter-killed male’s age, population density, and hunting intensity. Our results emphasize that hunting can have long-term indirect effects which can affect population viability
Journal Article