Sean D. Farley

Nitrogen and carbon isotope fractionation between mothers, neonates, and nursing offspring

Stable isotope signatures of lactating females and their nursing offspring were measured on 11 species, including herbivores, carnivores, hibernators, and non-hibernators. We hypothesized that: (1) nursing offspring would have stable isotope signatures that were a trophic level higher than their mothers, and (2) this pattern would be species-independent. The plasma of adult females had a δ15N enrichment over their diets of 4.1±0.7‰, but offspring plasma had a mean δ15N enrichment over maternal plasma of 0.9±0.8‰ and no C enrichment (0.0±0.6‰). The trophic level enrichment did not occur between mother and offspring because milk was depleted in both δ15N (1.0±0.5‰) and δ13C (2.1±0.9‰) relative to maternal plasma. Milk to offspring plasma enrichment was relatively small (δ15N enrichment of 1.9±0.7‰ and δ13C enrichment of 1.9±0.8‰) compared to the trophic level enrichment between the adults and their diets. While some species did have significant differences between the isotope signatures of mother and offspring, the differences were not related to whether they were hibernators or non-hibernators, carnivores or herbivores. Investigators wanting to use stable isotopes to quantify weaning or other lactation processes or diets of predators when both adults and nursing offspring are consumed must first establish the parameters that apply to a particular species/environment/diet combination.

Polar and brown bear genomes reveal ancient admixture and demographic footprints of past climate change

Polar bears (PBs) are superbly adapted to the extreme Arctic environment and have become emblematic of the threat to biodiversity from global climate change. Their divergence from the lower-latitude brown bear provides a textbook example of rapid evolution of distinct phenotypes. However, limited mitochondrial and nuclear DNA evidence conflicts in the timing of PB origin as well as placement of the species within versus sister to the brown bear lineage. We gathered extensive genomic sequence data from contemporary polar, brown, and American black bear samples, in addition to a 130,000- to 110,000-y old PB, to examine this problem from a genome-wide perspective. Nuclear DNA markers reflect a species tree consistent with expectation, showing polar and brown bears to be sister species. However, for the enigmatic brown bears native to Alaskas Alexander Archipelago, we estimate that not only their mitochondrial genome, but also 5–10% of their nuclear genome, is most closely related to PBs, indicating ancient admixture between the two species. Explicit admixture analyses are consistent with ancient splits among PBs, brown bears and black bears that were later followed by occasional admixture. We also provide paleodemographic estimates that suggest bear evolution has tracked key climate events, and that PB in particular experienced a prolonged and dramatic decline in its effective population size during the last ca. 500,000 years. We demonstrate that brown bears and PBs have had sufficiently independent evolutionary histories over the last 4–5 million years to leave imprints in the PB nuclear genome that likely are associated with ecological adaptation to the Arctic environment.

Sexual dimorphism, reproductive strategy, and human activities determine resource use by brown bears

Despite significant sexual dimorphism and differing reproductive strategies in carnivores, sexual segregation is rarely studied and is often overlooked in the management of wild populations. Potential nutritional constraints imposed by sexual dimorphism and differing reproductive strategies between the sexes have important implications, particularly when combined with differential effects of human activities on sex and age classes. We examined the effects of sexual dimorphism, reproductive strategies, and human activities (bear-viewing and hunting) on resource use by different sex and age classes of brown bears (Ursus arctos). Sexual segregation of habitat use and effects of experimental bear-viewing were quantified at a single site in south-central Alaska, U.S.A., by capturing, collaring, and observing brown bears at a salt marsh and salmon stream. Effects of salmon capture rate, availability of alternative salmon runs, harvest pressure, and numbers of annual visitors on sex and age class use were examined from data collected or previously published from 13 other sites. Bear-viewing sites on salmon streams where salmon capture rates were low (<4 salmon/hour) resulted in low use by adult males (<10% of all bears), except for sites with falls. However, maximum male use of viewing areas also depended on the availability of alternative salmon streams and harvest pressure. Use of habitats by females with dependent young was significantly related to the prevalence of adult males at the site. Thus, both sexual dimorphism and differing reproductive strategies led to sexual segregation in habitat use by bears. As a result of infanticide, females with young appear to prioritize avoidance of male bears over avoidance of humans when choosing habitats, in contrast to responses documented in herbivores. Because carnivores often exhibit both sexual dimorphism and infanticide, selection for sexual segregation is likely to be high. In these cases, the nutritional demands of large adult males, balanced with responses to human activity, drive dynamic temporal and spatial distributions of individuals in the population.

Predicting Body Condition of Bears via Two Field Methods

Bioelectrical impedance analysis (BIA) and isotopic water dilution have been proposed as methods suitable for determining the body composition of wild bears. We investigated the accuracy of these 2 techniques in a single-blind study. Water dilution was a more precise predictor of the body fat content of black bears (Ursus americanus) and brown bears (U. arctos) than was BIA, as indicated by comparison of the standard error of the estimate (SEE; 2.31% vs. 3.27%). The mean of the estimates from both methods predicted bear body fat content with the least error (SEE = 1.67%). The magnitude of the error for either method was not significantly related to body mass, lipid content, or animal age. Accurate, repeatable BIA estimates require extensive training and experience to standardize measurement conditions. Isotopic water dilution has fewer error sources but requires laboratory analyses.

Are inland wolf–ungulate systems influenced by marine subsidies of Pacific salmon?

Wolves (Canis lupus) in North America are considered obligate predators of ungulates with other food resources playing little role in wolf population dynamics or wolf prey relations. However, spawning Pacific salmon (Oncorhyncus spp.) are common throughout wolf range in northwestern North America and may provide a marine subsidy affecting inland wolf-ungulate food webs far from the coast. We conducted stable-isotope analyses for nitrogen and carbon to evaluate the contribution of salmon to diets of wolves in Denali National Park and Preserve, 1200 river-km from tidewater in interior Alaska, USA. We analyzed bone collagen from 73 wolves equipped with radio collars during 1986-2002 and evaluated estimates of salmon in their diets relative to the availability of salmon and ungulates within their home ranges. We compared wolf densities and ungulate:wolf ratios among regions with differing salmon and ungulate availability to assess subsidizing effects of salmon on these wolf-ungulate systems. Wolves in the northwestern flats of the study area had access to spawning salmon but low ungulate availability and consumed more salmon (17% +/- 7% [mean +/- SD]) than in upland regions, where ungulates were sixfold more abundant and wolves did or did not have salmon spawning areas within their home ranges (8% +/- 6% and 3% +/- 3%, respectively). Wolves were only 17% less abundant on the northwestern flats compared to the remainder of the study area, even though ungulate densities were 78% lower. We estimated that biomass from fall runs of chum (O. keta) and coho (O. kisutch) salmon on the northwestern flats was comparable to the ungulate biomass there, and the contribution of salmon to wolf diets was similar to estimates reported for coastal wolves in southeast Alaska. Given the ubiquitous consumption of salmon by wolves on the northwestern flats and the abundance of salmon there, we conclude that wolf numbers in this region were enhanced by the allochthonous subsidy provided by salmon and discuss implications for wolf-ungulate relations.

Dietary and spatial overlap between sympatric ursids relative to salmon use

Abstract We hypothesized that there would be minimal dietary overlap between sympatric brown bears (Ursus arctos) and American black bears (U. americanus) relative to salmon (Oncorhynchus spp.) utilization when alternative foods (e.g., fruits) are abundant. To maximize the chance that we would reject this hypothesis, we examined the diets of brown and black bears known to have visited salmon streams. Species, sex, and individual identification of bears visiting salmon streams were determined by DNA analysis of hair and feces collected in 2002–2004 along those streams. Diets were estimated from fecal residues and stable isotope analyses of hair. Assimilated diets of brown bears were 66.0% (SD  =  16.7%) salmon, 13.9% (SD  =  7.5%) terrestrial animal matter, and 20.1% (SD  =  17.2%) plant matter. Assimilated diets of black bears were 8.0% (SD  =  5.4%) salmon, 8.4% (SD  =  9.7%) terrestrial animal matter, and 83.6% (SD  =  7.7%) plant matter. Male and female brown bears did not differ in either the proportion of dietary salmon, terrestrial animal matter, or plant matter. The relative amounts of fruit residues in the feces of brown bears (87.0%, SD  =  15.2%) and black bears (91.8%, SD  =  7.2%) did not differ. Both sexes of brown bears visited salmon streams and consumed significant amounts of salmon, but only male American black bears visited streams and then consumed minimal amounts of salmon. Thus, brown bears were largely carnivorous and black bears were largely herbivorous and frugivorous. This reduced dietary overlap relative to salmon and fruit use is understandable in light of the concentrated, defendable nature of salmon in small streams, the widely dispersed, non-defendable nature of abundant fruits, the dominance of brown over black bears, the higher energy requirement of the larger brown bear, and, therefore, the differing ability of the species to efficiently exploit different food resources.

Nutritional consequences of experimentally introduced tourism in brown bears.

Abstract Although numerous studies have documented behavioral effects of nature-based tourism on wildlife populations, few studies have determined whether behavioral changes translate to effects on individual condition and population health. This issue is currently a concern for wildlife managers in Alaska, USA, and Canada where bear viewing is a rapidly growing industry expanding into previously undisturbed bear habitats. Rather than record observations at long established tourism sites, we experimentally introduced bear viewing into 2 relatively undisturbed brown bear (Ursus arctos) populations in south-central Alaska. We examined the nutritional consequences of behavioral changes induced by the presence and activity of bear viewers for bears feeding on early summer vegetation and late-summer salmon (Oncorhynchus kisutch and O. nerka). We used Global Positioning System collars, monitored food resource availability, and quantified individual resource use and condition for a year prior to and during the introduction of bear viewing. Though bear viewing altered spatiotemporal resource use in all treatments, total resource use declined only when we exposed bears to 24-hour daily human activity. Energy expenditure, indexed as daily travel distances, was significantly higher when bears responded by altering spatial rather than temporal resource use. However, body weight and composition were unaffected by all treatments as bears shifted their foraging to other locations or times. Managers can minimize nutritional impacts of bear-viewing programs by avoiding spatial displacement and providing predictable time periods when bears can access food resources free of human activity. Bears in this study exhibited a high degree of behavioral plasticity, which may be an important factor in identifying flagship species for sustainable ecotourism programs.

Importance of salmon to wildlife: Implications for integrated management

Abstract Salmon (Oncorhynchus spp.) are an important resource for terrestrial wildlife. However, the salmon requirements of wildlife populations and the role wildlife play in nutrient transport across ecosystems are largely ignored in salmon and habitat management. Any activity that reduces the availability of or access to salmon by wildlife may adversely affect wildlife populations and, potentially, ecosystem-level processes. Thus, when the conservation of specific wildlife populations or healthy ecosystems is the management objective, allocation of salmon to wildlife should be considered. We provide an example of how such allocations could be calculated for a hypothetical bear population. Ultimately, salmon allocation for wildlife calls for integrated management of natural resources across agencies, across species, and across ecosystems. We summarize the current state of knowledge relative to the interaction between Pacific salmon and the terrestrial ecosystem, with special emphasis on the import of salmon to terrestrial wildlife and the import of wildlife to terrestrial and aquatic ecosystems.

Patterns of landscape use by female brown bears on the Kenai Peninsula, Alaska.

Abstract We describe landscape use of female brown bears (Ursus arctos) on the Kenai Peninsula, Alaska, USA. Radiocollars, fitted to 43 adult female brown bears, provided radio relocations, which we used to describe habitat use patterns by season and reproductive class at the landscape scale. Brown bears were associated with areas with low densities of human developments and roads, as well as riparian areas that were close to cover. Presence of streams and lakes that supported spawning salmon (Oncorhynchus spp.) positively influenced summertime distribution of bears. Female brown bears with cubs avoided concentrations of other bears. Resource managers may use this information to respond to brown bear conservation issues associated with increasing human populations and associated development in the establishment of road density standards, seasonal road closures, management of recreation sites, and vegetation management on the Kenai Peninsula.

Genetic characterization of Kenai brown bears (Ursus arctos): Microsatellite and mitochondrial DNA control region variation in brown bears of the Kenai Peninsula, south central Alaska

We collected data from 20 biparentally inherited microsatellite loci, and nucleotide sequence from the maternally inherited mitochondrial DNA (mtDNA) control region, to determine levels of genetic variation of the brown bears (Ursus arctos L., 1758) of the Kenai Peninsula, south central Alaska. Nuclear genetic variation was similar to that observed in other Alaskan peninsular populations. We detected no significant inbreeding and found no evidence of population substructuring on the Kenai Peninsula. We observed a genetic signature of a bottleneck under the infinite alleles model (IAM), but not under the stepwise mutation model (SMM) or the two-phase model (TPM) of microsatellite mutation. Kenai brown bears have lower levels of mtDNA haplotypic diversity relative to most other brown bear populations in Alaska.