Ecosystems are linked by spatial subsidies, the bi-directional flows of nutrients, materials and energy that cross ecosystem boundaries. Considered one of the planet’s most productive and diverse meta-ecosystems, the broad interface between land and sea is crossed by innumerable abiotic and biotic spatial subsidies, including migratory animals. Routinely crossing ecological boundaries, migrants play significant roles in subsidizing receiving ecosystems, including influencing ecosystem productivity, diversity, community structure and trophic cascades. On the Pacific coast of North America, spatial subsidies driven by migratory Pacific salmon have been intensively studied. Like many of the world’s migrants, however, salmon populations have declined considerably and most of our scientific knowledge has been gained from a diminished subsidy. Other subsidies, including those driven by migratory species in decline, remain relatively unknown. Each year, Pacific herring (Clupea pallasii) migrate to shallow waters to spawn on nearshore and intertidal substrates. Despite suggestions in the literature that herring, an abundant, nearshore/intertidal spawning forage fish, subsidizes coastal ecosystems, there had been no investigation of cross-ecosystem interactions. Just as stable isotopes and fatty acids have been used to explore wrack (drift macrophytes) subsidies to intertidal ecosystems, we combined both approaches to trace the input of Pacific herring and wrack to semi-terrestrial amphipods (Traskorchestia spp.), which are highly abundant detritivores in beach ecosystems. Brown algae and seagrass were major contributors to amphipods but when available, herring was also a significant resource. Because amphipods are prey for terrestrial consumers, including bears (Ursus spp.), we also identified indirect trophic linkages between herring and terrestrial ecosystems. Bears are major consumers and vectors of salmon into terrestrial ecosystems, but little is known regarding their involvement in other spatial subsidies. Using a model-based inference approach paired with remote cameras to monitor intertidal black bear (U. americanus) activity, we determined that the best predictors of black bear intertidal activity were major intertidal prey items (herring and amphipod biomass) and Julian day. Bears positively responded to herring and amphipod biomass on beaches but it was the analysis of scats that determined the contribution of herring eggs to the diets of bears. In 2010, the herring spawn was relatively poor and consumption of eggs was negligible, with amphipods constituting a major portion of bear diets. In following years, herring egg loading was relatively high and eggs were the dominant dietary item in bear scats. Tracing the contribution of herring into terrestrial areas proved challenging and instead, we furthered knowledge of the within-watershed spatiotemporal influences of salmon on conifer tree ring growth and δ15N signatures. Both tree ring growth and δ15N signatures tracked the known spatial distribution of salmon carcasses. Using a model-based inference approach, salmon abundance and interaction terms of salmon*temperature and salmon*distance into the forest best predicted tree growth. In contrast, salmon abundance was not a leading predictor of δ15N. By broadening our understanding of the fine-scale influence of salmon on a stand of ancient trees, this research is expected to contribute to future exploration of the terrestrial influences of Pacific herring. / Graduate / 0329 / cfox@uvic.ca

Thesis (M.A.)--Western Washington University, 2009. / Title from PDF title page (viewed on June 29, 2009). Includes bibliographical references.

Thesis (M.S.)--University of Tennessee, Knoxville, 2009. / Title from title page screen (viewed on Nov. 3, 2009). Thesis advisor: Joseph Clark. Vita. Includes bibliographical references.

Using specially trained scat detection dogs we located fecal samples from black bear (Ursus americanus) and coyote (Canis latrans) throughout three study areas in Newfoundland, Canada, to describe these predators diet. Our sampling efforts were designed around seasons which were important to woodland caribou (Rangifer tarandus caribou) calving and resource use. We identified hairs microscopically to prey species and grouped other remains to facilitate our analysis. Bear exhibited an omnivorous diet throughout the study areas, ecological seasons and inside and outside the caribou calving grounds while coyote were limited to caribou, moose and snowshoe hare.

Individual variation and fitness are the cornerstones of evolution by natural selection. The trophic niche represents an important source of phenotypic variation on which natural selection can act. Although individual variation is fundamental to species-level ecological and evolutionary change, individual variation is often ignored in population-level approaches to wildlife ecology, conservation and management. Failing to link individual resource use to fitness or to biological outcomes related to fitness limits us to managing for the average resource needs of a population, which may be insufficient for protecting the diversity of resource use within populations and the underlying eco-evolutionary processes that generate that diversity. My goals were to provide insights into the mechanisms that generate and constrain intrapopulation trophic niche variation, evaluate whether linkages exist between individual biological outcomes and variation in food habits across the range of resources consumed within generalist consumer populations and examine how that variation manifests in population-level responses. I investigated the causes and physiological consequences of intrapopulation trophic niche variation in two generalist consumers, the American black bear (Ursus americanus) and brown bear (U. arctos) across three sites in British Columbia, CAN and at one site in Alaska, USA. My primary tools included stable isotope analysis to estimate diet, enzyme-linked immunoassay of hair to quantify the hormone cortisol for indexing physiological stress, and genetic analyses to identify individuals, species, and sex and to estimate ancestry. I found that individual differences in resource use can result in similar biological outcomes and that similar resource use can result in different biological outcomes. Intra- and interspecific competition, sex-based differences in nutritional and social constraints and annual variation in food availability all influenced trophic niche variation and the resultant biological outcomes. I also found evidence of a link between intrapopulation trophic niche variation and population genetic structure. My results highlight the diverse ecological drivers and diverse consequences of trophic niche variation, which further illuminates why the trophic niche is a nexus for eco-evolutionary dynamics.

I monitored cause-specific mortality and factors influencing mortality risk for white-tailed deer in the Upper Peninsula of Michigan, USA, during two high mortality risk periods: adult female deer during Feb–May, and fawns from birth to 6 months. I observed high rates of predation and starvation for adult female deer during Apr–May, suggesting that late winter represents a survival bottleneck due to nutritional declines. A strong negative relationship existed between snow free days during late winter and mortality risk. Predation was the dominant mortality source for fawns but predation risk decreased with larger birth mass. Black bears and coyotes accounted for most fawn kills at the population level, but wolves and bobcats had greatest per-individual fawn kill rates. My results suggest predation was the dominant mortality source for fawns and adult female deer, but multiple predator species were important and nutritional condition of deer influenced their vulnerability to predation.

Understanding how wildlife utilize habitat at varying scales is important for understanding and predicting potential impacts of landscape changes (e.g., habitat loss and fragmentation, restoration efforts, climate change, etc.) and in determining effective strategies for conservation and management. This research examines fine-scale and landscape-level habitat use of black bears in Canyon de Chelly National Monument (CACH), Arizona, USA in the context of large-scale landscape change. Currently, CACH is undergoing a large-scale restoration effort to remove all of the non-native Russian olive (Elaeagnus angustifolia) and tamarisk (Tamarix ramosissima and T. chinensis) within the monument. As black bears rely on the Russian olive as a primary food source, a major goal of this research was to evaluate how bears currently use this resource and how its removal might affect bear habitat use and bear-human interactions within the monument. I developed a model of 3rd order (fine-scale) black bear habitat use in CACH using an occupancy modeling approach. Model results indicated that fine-scale habitat selection by bears in CACH is being driven by the presence of non-native Russian olive as well as native food sources. Thus availability of native foods may be sufficient to maintain the bear population in CACH, and bears may quickly adapt to the loss of the non-native food source. Similarly, results showed that bears avoid human areas and farmsteads and prefer higher elevations, suggesting that once olive is removed in the lower canyons near human areas, bear-human interactions may become less of a problem over the long term. I also developed a model of 2nd order (landscape-level) habitat use and evaluated movement patterns of black bears in CACH using location data collected from GPS collars. Model results showed that bears selected areas with higher tree canopy cover and terrain ruggedness, indicating that forest cover and escape cover are primary factors driving black bear habitat selection at the landscape scale in this region. Movement patterns revealed large mean daily movements and low average turning angles, indicating long, linear movements designed to take advantage of the mosaic of available habitats and food resources available over larger areas. I extracted DNA from hair samples collected throughout the study area to examine genetic variability and population structure of black bears in the region. Analyses revealed a relatively healthy, panmictic population across the wider landscape. No substantial genetic structuring was observed in multiple analyses, though I did find evidence of a slight isolation-by-distance pattern within the population. Measures of both current (Nb = 24) and long-term (Ne = 579) effective population size indicated a relatively high number of breeders in the current population and a sufficient amount of gene flow within the larger "superpopulation" to maintain long-term genetic viability. I focused the final portion of my dissertation research on understanding the factors that influence stakeholder acceptance capacity for black bears, which plays a central role in contemporary wildlife management issues, including human-wildlife conflict. I used data from telephone interviews of 1,546 residents in Virginia to develop conceptual models of black bear acceptance capacity at both county and state levels. Model results suggested that more deep-seated and less easily influenced factors (e.g., values and risk perceptions) are at the heart of stakeholder attitudes and perceptions of wildlife, making influencing these perceptions more difficult. Agencies can indirectly affect these attitudes and perceptions, however, by targeting more potentially pliable factors related to wildlife acceptance capacity such as knowledge, personal experience with wildlife, and trust in management agencies. / Ph. D.

The effects of gypsy moth infestation on the Shenandoah National Park (SNP) black bear population and habitat were studied during 1985 - 1991 by comparing radio telemetry, population, and behavioral data from preinfestation years (1982 - 1986) and years with extensive defoliation (1987 - 1991). Gypsy moth defoliation (> 60% canopy loss) increased from 546 ha in 1986 (1 % of the study area), to 2,304 ha in 1987 (4%), 6,227 ha in 1988 (12%), and 17,736 ha in 1989 (34%). Chestnut oak and red oak habitat types received the greatest defoliation; 60% and 45% of these habitat types suffered greater than 60 % canopy loss in the North and Central Districts, respectively. Infestation resulted in a 99% reduction in acorn production in defoliated stands. Maximum daily temperatures 0.5 m above the ground in defoliated stands averaged 4.7 ± 0.3 C, 4.3 ± 0.4 C, and 2.5 ± 0.3 C warmer (P < 0.01) than in nondefoliated stands during peak defoliation, refoliation, and post-refoliation periods, respectively. Bear / Ph. D.

The Cooperative Alleghany Bear Study (CABS) was initiated in 1994 to address concerns over the lack of biological and ecological data for black bear (Ursus americanus) populations in the Alleghany Mountains of western Virginia. I examined home range dynamics of bears during 1994-2002 on 2 study areas that were approximately 160 km apart. I analyzed my data with 3 home range programs (AMA, HRE, and ABODE) and determined the HRE was the least biased and produced the most biologically reasonable home range estimates. I used HRE to generate annual home ranges (fixed-kernel) for 90 bears over 160 bear years; I also generated seasonal home ranges using MCP. Annual and seasonal home ranges of male and female adult bears in the southern study area were larger than that of male and female adult bears in the northern study area, respectively; southern females and northern males had annual home ranges similar in size at the 95% and 75% fixed-kernel contours. In both study areas, most bears did not shift their range when transitioning from spring to summer (North: 63.0%; South: 57.0%) or from summer to fall (North: 67.0%; South: 65.0%), while most bears shifted their seasonal range between spring and fall (North: 67.0%; South: 52.0%). Most female bears in both study areas maintained the same spring and summer home range throughout the duration of the study, while 63% of northern females changed their fall home range and 55% of southern females maintained their fall home range. I found no differences in annual and seasonal home range size among years or among age classes for adult females, but tests for intra-year seasonal difference indicated that fall range was larger than spring and summer in 1997, when western Virginia experienced a poor mast crop. Females with and without COY had similar annual home ranges in either study area. In the north, seasonal home range size did not differ between females with and without COY, while in the south, breeding females (i.e. without COY) had larger spring ranges and smaller fall ranges than females with COY. In both study areas, females with COY had larger fall home ranges than during spring, while seasonal ranges of breeding females did not vary in size during the year. / Master of Science

Following mass agricultural expansion and deforestation in the late 1800s and early 1900s, American black bear populations (Ursus americanus, hereafter black bear) were low across most of Virginia. The Department of Wildlife Resources (VADWR) and the United States Forest Service worked to rebuild wildlife habitat in the state, leading to recovery of bears. While some aspects of black bear hibernation and foraging ecology have been studied, there remain knowledge gaps on key aspects of black bear biology. Hibernation behavior, for example, is understudied with most knowledge stemming from post-hibernation emergence studies. We used a unique dataset with continuous monitoring of mother bears and their cubs from Virginia Tech's Black Bear Research Center in 2015-2016, to better understand hibernation ecology, mother-cub dynamics, and biological-foster cub dynamics. We found that increased black bear activity levels were driven by both higher temperature and photoperiod, but the onset of hibernation was primarily driven by only temperature. Temperature is projected to rise in VA and rising temperatures regionally could be responsible for increased active behaviors in bears, which could lead to increased human-black bear interactions on the landscape, especially during hibernation onset. We also found that mother bears spend similar amounts of time with biological and foster cubs. Biological cubs did not show more dominance behaviors toward fosters than biological siblings, which we also observed with foster, where they did not continuously display these behaviors towards biological cubs. These results are promising for orphan cub fostering programs in VA and other states and indicate that foster cub litter integration can be successful. Foraging ecology also is challenging to study in black bears because direct observation is generally not possible due to their cryptic nature, closed habitat, or potential danger in observing. We used a dataset from camera collars deployed on 15 bears (8 males, 7 females) in Bath County Virginia in 2018 and 2019 to better understand diet seasonality and to determine habitat and environmental drivers of black bear foraging patterns, particularly on white-tailed deer (Odocoileus virginianus) and invasive plant species. We identified 178 unique diet items to family, genus, or species in videos, much higher numbers than previously reported in the literature, and we found high diet overlap between sexes. Diet composition was primarily influenced by season with higher levels of consumption of herbaceous soft mast in spring, fruit and seed soft mast in summer, and hard mast in fall. Females exhibited more hunting events on deer fawns in spring than males, but males and females consumed similar numbers (28 vs. 24, respectively) via hunting and scavenging combined. Males consumed anthropogenic foods more often than females, particularly when closer to human settlements and males more commonly consumed invasive plant species in spring while females more often consumed insect in spring invasive species in summer. Our results highlight strong seasonally- and subtle sex-mediated differences in black bear diets. We provide information on drivers of diet choices by bears, as well as identify where foraging hotspots on species of interest occur, providing information useful to VADWR in managing increased human-wildlife interactions (and mitigating potential for negative interactions), predator-prey relations, and invasive species spread across the landscape. / Master of Science / American black bears were almost eliminated from Virginia following mass agricultural expansion and deforestation in the late 1800s and early 1900s. The Department of Wildlife Resources (VADWR) and the United States Forest Service worked to rebuild wildlife habitat in the state, leading to recent recovery of bears in VA. However, there are still gaps in our knowledge of black bear ecology and gaining new knowledge will aid in better management of black bears across the state. We used two unique video data sets to study hibernation and foraging (or feeding) ecology to fill important knowledge gaps. To study hibernation behavior, we used a video data set of continuously monitored, temporarily captive mother bears and their cubs held at Virginia Tech's Black Bear Research Center in 2015-2016. We found that black bear activity was driven by both daily temperature and photoperiod (length of daylight), but onset of hibernation was mostly driven by temperature only. With Virginia projected to experience rising, and more variable, temperatures due to climate change, this may mean that bears will stay longer on the landscape in fall, even past the point where vegetation (food) has gone dormant, potentially leading to increased human-black bear interactions as bears seek human foods. We also examined mother-cub interactions and found mother bears did not spend more time with their biological cubs than their fosters (fosters are orphaned black bear cubs arising due to death of mothers) and biological cubs did not show more dominance behaviors (i.e., biting, chasing, or pinning to ground) toward fosters than biological siblings. These are positive results for VADWR's surrogate sow program across the state. Foraging ecology is difficult to study in black bears because direct observation is generally not possible due to their secretive nature, dense habitat, or potential danger in observing. Therefore, traditionally, many studies collect and examine scat (fecal) samples or stomach contents from harvested bears, but these methods may not find diet items that are quickly digested. We used a unique dataset from video camera collars placed on 15 bears in Bath County Virginia in 2018 and 2019. We found some important differences from the literature - our spring diets had higher proportions of deer and soft vegetation (leaves), however the unknown diet items in the literature were high, leading to incomplete or incorrect diet profiles. Diet composition was primarily influenced by season with higher levels of consumption of leafy soft mast in spring, fruit and seed soft mast in summer, and hard mast in fall. Males and females had high dietary overlap and we found that females had more hunting events on deer fawns in spring than males, but overall males and females consumed similar numbers (28 vs. 24, respectively) via hunting and scavenging combined. Males more often consumed human foods than females, while females more often consumed insect species than males. Also, males more commonly consumed invasive species in spring, while females did so in summer. Our results highlight differences in diet content caused by season and subtle differences in diets between sexes. We provide information on drivers of feeding choices bears make and we identify foraging hotspots for deer and invasive species, information useful to VADWR in managing potential human-wildlife interactions, predator-prey relations, and invasive species spread across the landscape.