Cole Burton

Understanding the complex mechanisms driving wildlife declines is crucial for effective conservation. Inthis thesis, I used camera traps to investigate two key conservation questions involving interspeciesinteractions in the Chilcotin Plateau of west-central British Columbia, Canada: 1) the role of disturbance-mediated apparent competition (DMAC) in the decline of the Itcha-Ilgachuz caribou population, and 2)the impacts of feral horses on native ungulate species.Caribou populations in Canada have faced significant declines, with DMAC emerging as an importantcontributor. I examined the mechanism of decline for the Itcha-Ilgachuz caribou, focusing on post-disturbance productivity, primary prey responses to disturbance, predator responses to prey anddisturbance, and caribou overlap with predators and disturbance. I found a productivity pulse followingdisturbance, but low overall productivity. Primary prey exhibited strong preferences for burnt areas, butnot cutblocks. Wolves were associated with primary prey, but not disturbances, and other predatorsshowed associations with primary prey and/or disturbances. Caribou did not avoid cutblocks, potentiallyincreasing their vulnerability to non-wolf predators. These findings provide mixed support for DMAC,and suggest that wildfire may be an important contributor to caribou decline in this population,emphasizing the importance of fire management going forward.I investigated the potential for competition between feral horses and two important native ungulates:moose and mule deer. Despite concerns about competition for wetlands and water resources, I foundno strong evidence of competitive exclusion or avoidance by moose or mule deer in the presence offeral horses. While moose and feral horses shared a preference for wetlands, none of the species hadincreased use of areas near water sources. Although mule deer showed decreased use in areasoverlapping with horses, this pattern was not conclusively linked to horse presence. These resultssuggest that feral horses may not be significantly impacting native ungulates in this region throughcompetition, but further research is warranted to explore other impacts such as grazing and apparentcompetition.This research underscores the complexity and context-dependency of conservation challenges involvinginteracting species, emphasizing the need for management strategies informed by population-specificknowledge to effectively conserve threatened species and ecosystems.

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Monitoring animals in northern Canada is especially difficult, and there is a need for cost-effective methods and sampling designs. Camera traps and autonomous recording units (ARUs) are both promising tools, but they are rarely combined. This thesis focuses on two areas of integration: sampling design and habitat modeling. Data were collected in Ts’udé Nilįné Tuyeta, an Indigenous and Territorial Protected Area in the Northwest Territories.Hierarchical sampling designs, where sensors are deployed in clusters, are common for ARUs but rarely used for camera traps. I evaluated a hierarchical design for camera traps using resampling to determine the sample size required to estimate detection rate, habitat associations, and species richness across our study area. I found that 4 cameras per cluster across 35 clusters were sufficient for most of the metrics tested, but up to 13 cameras per cluster were needed to precisely estimate detection rate of rarer species. Cluster-scale species richness was unreliable even with up to 18 cameras per cluster, suggesting richness is best estimated at a larger scale.In my third chapter, I evaluated the use of ARUs and camera traps to create habitat models for sandhill crane Antigone canadensis. I tested for differences in the spatial scale of inference by performing model selection using two buffer sizes, and whether a model that integrated both data types would improve predictive performance. I found that ARUs made the best inferences at the landscape scale, while cameras made the best inferences at the home range scale. The integrated model did show evidence for hierarchical habitat selection by cranes, but did not improve predictive performance. Overall, I found that each data source had merits of its own, but that, in this case, integration into one model was not an improvement over modeling data separately.These two data chapters showcase two benefits of pairing camera traps and ARUs: first, improving response variables by carefully considering sampling design; and second, improving our understanding of habitat selection by using data from both sensors. Efforts should continue to apply and test these and other methods for effective wildlife monitoring in Canada’s rapidly changing northern environments.

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Woodland caribou are a threatened species in Alberta and across Canada, due to anthropogenic disturbances, such as forest harvest and linear features, which facilitate increased predation on caribou. To aid caribou recovery, the Government of Alberta is working to conserve areas of existing caribou habitat, recover habitat on linear features, and annually reduce wolf abundance. These management actions are beneficial for caribou, but potential effects on other wildlife have rarely been tested. To better understand any effects, we conducted multispecies surveys using remote cameras (n=60) within and surrounding the Little Smoky caribou range in west-central Alberta.We hypothesized that changes in wolf occurrence rate would be the predominant factor influencing other wildlife, because they are the top predator on the landscape and have been shown to exert top-down control in other ecosystems. We predicted decreased wolf occurrence rate in areas of high wolf removal efforts would result in higher occurrences of coyotes and lynx, through decreased competition, and higher occurrences of moose, elk, and deer, through decreased predation. Our alternative hypothesis was that habitat disturbances would be the dominant factor determining species occurrences through bottom-up ecosystem control. We used GLMMs to test the effects of anthropogenic disturbance and wolf population reduction on several medium and large-bodied mammal species.As expected, wolf occurrence was negatively affected by wolf removals. Wolf population reduction reduced wolf occurrences annually, but wolves recolonized areas of high removal effort each year. Unexpectedly, mesopredator occurrences were positively associated with, and ungulate occurrences unaffected by, rates of wolf occurrence. These species were instead more strongly associated with anthropogenic habitat disturbances (forest harvest, linear features), and mesopredators occurrences were also associated with prey availability. Our results suggest that despite the direct effect of wolf removals on wolves, wolf population management did not have cascading effects on other wildlife in this system. Rather, bottom-up factors, such as habitat features and prey availability were the most important drivers affecting the medium- and large-bodied mammal community in west-central Alberta.

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Active forest management for timber production, through the harvesting of forest stands using cut blocks, frequently overlaps grizzly bear (Ursus arctos) habitat on multi-use landscapes in North America. Thus, it is critical to understand how forest harvest management can effectively support grizzly bear conservation efforts. While many localized studies have investigated relationships between forest harvest and grizzly bear habitat use, a synthesis of our current understanding of these complex interactions is warranted. This thesis reviewed publications that empirically assessed grizzly bear use of recently harvested forest stands (
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Globally, protected areas face a challenge of meeting the dual mandate of protecting biodiversity while providing recreational enjoyment for humans. Balancing these mandates is confounded by limited information on species status and insufficient recreation monitoring. Effective protected area (PA) management is critical for wildlife in this time of pervasive human impact known as the Anthropocene. Where non-consumptive human recreation is considered low impact, with growth in outdoor recreation it is important to know if recreation is impacting wildlife and how. Using camera traps, I assessed the potential for recreational impacts on mammal habitat use in space and time, in Cathedral Provincial Park, British Columbia, Canada. I also estimated population density for an at-risk mountain goat (Oreamnos americanus) population using two methods: spatial capture-recapture (SCR) and spatial mark-resight (SMR). I assessed recreational impacts on habitat use at the weekly scale, while also evaluating daily activity patterns for eight mammal species. I hypothesized that coexistence with recreation would require spatial and/or temporal niche segregation, with large carnivores being most sensitive and exhibiting spatial avoidance as suggested by the predator shield hypothesis. I predicted that mesocarnivores and ungulates would exploit this “shield” spatially, while exhibiting temporal avoidance of humans. I found spatial co-occurrence between ungulates and recreation, suggesting that these species may be using people as a shield from predators or for nutritional subsidies, but did not see the predicted negative relationship between predators and humans, except for coyotes (Canis latrans). Temporally, all species other than cougars (Puma concolor) had activity patterns significantly different from that of recreationists, suggesting stronger displacement in the temporal niche, while wolves (Canis lupus) and mountain goats showed significantly different use of on and off-trail habitat in time.Estimates of mountain goat density varied between methods, from a minimum 6.32 (95% CI; 2.98-13.40) to a maximum of 11.54 (6.97-19.13) goats per 100 km2. I found SMR estimates to have higher precision than SCR estimates across all three years. With this study I show that camera trap surveys can be used to assess interactions between wildlife and recreation, while also providing basis for monitoring population trends in sensitive species.

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The dual-mandate for protected areas (PAs) to simultaneously promote recreation and conserve biodiversity may be hampered by negative effects of recreation on wildlife. However, reports of these effects are inconsistent, presenting a knowledge gap that hinders evidence-based decision making. To narrow this gap, I used camera traps to monitor human activity and terrestrial mammals in an exurban PA and an adjacent research forest, with the objective of discerning impacts of human activity on the habitat use and diel activity patterns of cougars (Puma concolor), black bears (Ursus americanus), black-tailed deer (Odocoileus hemionus), snowshoe hares (Lepus americanus), coyotes (Canis latrans), and bobcats (Lynx rufus). I then used Bayesian models to assess how wildlife and humans share space and time. First, I investigated whether recreationists and motorized vehicles displaced these focal species at the weekly temporal scale. I found that hikers negatively affected bobcat habitat use, whereas vehicles negatively affected black bear habitat use. I then explored how species’ detection rates shifted during an unexpected period of park closure (due to COVID-19). I found increased cougar detection rates when public access was restricted, but subsequent decreases in cougar detection rates and increases in black-tailed deer detection rates upon the park’s reopening. Finally, I investigated how these species’ diel activity patterns were impacted by human pressures. Only black bears shifted to be (moderately) more nocturnal in spaces or times of higher observed human activity, whereas coyotes, snowshoe hares, cougars, and black-tailed deer all altered their diel activities in response to landscape features associated with human activity (e.g., trail or road densities). My results illustrate that wildlife may be displaced by human activity, but this displacement is often species- or activity-dependent and occurs at various spatiotemporal scales. I also provided support for the use of camera traps to simultaneously monitor human and wildlife activities and encourage PA managers to consider whether recreation is negatively impacting conservation goals within their own PAs. I stress that recreation has the potential to offset PA natural resource conservation goals. However, further research is needed to understand the how displacement by recreation might translate to consequences for wildlife populations.

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British Columbia’s interior forests have been heavily logged, burnt and subject to beetle outbreaks for decades. The compounding effects of these disturbances on wildlife and their habitat must be considered. Partial retention forest harvesting may be a method that could mitigate some of the negative effects of clearcut harvesting on wildlife. However, tests of the effects of partial harvests on ecosystem patterns and processes in different contexts are needed. From December 2018 and June 2020, we conducted live trapping for small mammals and camera trapping for medium-to-large-bodied mammals to estimate species diversity, population density, habitat use, and behaviours across different forest harvesting practices across a 900 km gradient in John Prince Research Forest, Alex Fraser Research Forest, and Jaffray (east Kootenays), BC. We detected 7 small mammal species, with diversity highest in the control (mean Shannon Index = 1.01, SE = 0.14) and partial retention treatments (means = 0.99, 0.98; SE = 0.17, 0.17) and significantly lower in the seed tree treatment (mean = 0.63, standard error = 0.17, p-value = 0.02). Population densities of North American deer mouse (Peromyscus maniculatus) and Southern red-backed vole (Myodes gapperi) estimated with spatially explicit capture-recapture models highlight the importance of partial harvest practices that maintain sufficient cover to support a higher diversity of small mammals and higher densities of forest specialists. Our medium- to large-bodied mammal diversity analysis suggests that the regional environmental context had a stronger effect on mammal communities than local-scale differences in harvesting practices. Vegetation productivity measured with normalized difference vegetation index was a more important predictor of habitat use for ungulates than harvest treatment, potentially due to the importance of forage availability. Across both small and large mammals, responses to forest harvesting were variable; several species used partial harvests more than clearcuts. Forest practices should consider broader implementation of partial harvests to provide suitable habitats for a broader range of species. More experimental approaches to forest operations are needed across larger spatial scales, such as adaptive management of forest practices with rigorous wildlife monitoring to ensure ecological objectives are met.

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The Andean bear (Tremarctos ornatus) is the largest carnivore in the tropical Andes and is an essential apex predator. Andean bears are vulnerable to extinction, and human-caused disturbances are driving population declines. Although the presence of free-ranging cattle is a major disturbance within protected areas, the effect of cattle on Andean bears is poorly understood. We used camera traps and remotely sensed data to assess the spatial and temporal relationships between cattle and bears in a protected area in northern Peru from 2015 to 2016. We hypothesized that cattle grazing represented a disturbance for Andean bears and predicted that bears would avoid cattle in space and/or time. We further predicted that the effects of cattle would be stronger during the wet season when their abundance and activity was higher. We tested the spatial prediction using generalized linear models, where we expected a negative relationship between the occurrence of cattle and bears. We included other factors potentially influencing bear occurrence in our models, including other measures of anthropogenic disturbance (occurrence of humans and dogs, and proximity to towns and farms) and of natural habitat variation in the refuge (elevation, slope, and forest cover). To test for temporal avoidance, we estimated the degree of overlap between daily activity patterns of bears and cattle. As predicted, we found a negative spatial association between bears and cattle and bears and humans and dogs. Bears were also less likely to occur closer to towns and farms adjacent to the refuge. Overall, bear responses to anthropogenic disturbance were stronger than to natural habitat variation. Surprisingly, the spatial avoidance of cattle by bears was stronger during the dry season. We did not find evidence of temporal avoidance, as there was high overlap between the daily activity patterns of bears and cattle, and between bears and humans and dogs, suggesting the potential for interaction where they do spatially co-occur. Given the threatened status of Andean bears, and the critical role of protected areas in their conservation, we recommend effective management of cattle and associated disturbances to protect and recover populations of this ecologically and culturally important carnivore.

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The urban-wildland interface is growing as human development expands, potentially increasing human-wildlife conflict. Conflicts include animals accessing garbage, damaging agricultural crops, or depredating livestock. For mammalian carnivores this often leads to lethal mitigation. Mortality from conflict represents a major threat to carnivores who miscalculate the risk of human-dominated areas. By contrast, carnivores that adapt to these novel anthropogenic environments may facilitate human-wildlife coexistence. Human-carnivore conflict is an increasing issue on Vancouver Island, British Columbia, due to rapidly expanding development and high concentrations of black bears (Ursus americanus) and cougars (Puma concolor). To reduce these conflicts and promote coexistence, it is critical to target proactive mitigations using reliable evidence to distinguish where conflict is probable from where carnivores are adapting to coexist. I modelled relative conflict probability using seven years of reported conflicts and GIS data to investigate which anthropogenic and environmental predictors best explained the spatial and temporal distribution of conflict in Victoria’s Capital Regional District. I found that the probability of conflict for both species increased along the urban-wildland interface, where human disturbance adjoined natural habitat. Black bear conflict also increased in rural areas in autumn before winter denning. I subsequently used a camera trap survey to see when and where bears were active across a gradient of human disturbance and compared bear habitat use to the previously estimated probabilities of conflict. For much of the year, bears used areas of low to medium conflict, such as forests near urban areas, avoided areas of higher human density, and were more nocturnal in urban and rural areas compared to wild. However, in autumn, bears were more active in areas of high conflict probability, specifically rural lands with ripe crops. This suggests that bear behaviour may allow for coexistence in most seasons by spatially and temporally avoiding humans, except in autumn when hyperphagia and peak anthropogenic crop availability increase the risk of human-bear conflict.Overall, I recommend proactive conflict mitigation to secure anthropogenic attractants against multiple carnivore species, and a particular focus on mitigations during seasonal peaks in attractive human food resources.

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A leading cause of large carnivore declines is conflict with humans, specifically due to livestock depredation. This conflict threatens both carnivore populations and human communities with livestock-dependent livelihoods. The expanding dairy industry in Sri Lanka, home to the endangered Sri Lankan leopard (Panthera pardus kotiya), provides an opportunity for proactive conflict mitigation. Little is known about incidents of livestock depredation or attitudes of pastoralists whose livelihoods may be threatened by leopard conflict. This thesis aimed to combine social and ecological research methods to address these knowledge gaps. We surveyed two pastoralist communities that differed in leopard conflict and socioecological factors (Yala and Central Hills) to identify determinants of attitudes towards leopards. We conducted Exploratory Factor Analysis and ran generalised linear models (GLMs) to detect influential variables. In the higher conflict region (Yala), attitudes towards leopards were positively related to socio-demographics (age, number of dependents, years rearing livestock) and an overall desire for wildlife conservation, while attitudes were negatively related to general awareness of leopard ecology and leopard-related tourism. In the lower conflict region (Central Hills), attitudes were positively related to a desire for increased government assistance in cattle rearing. The inability to own land were common concerns for pastoralists in both regions. We recommend assessing programs that may improve attitudes towards leopards, such as involving pastoralists in tourism programs and restricted land ownership. While the Central Hills is currently not experiencing depredation, proactively addressing hardships (e.g. improving roads, subsidizing cattle feed) may facilitate positive attitudes, should incidents of conflict increase. To investigate potential drivers of cattle depredation in Yala, we used GLMs to test the importance of hypothesised explanatory variables, specifically: native prey availability, cattle husbandry, number of cattle, distance from national park, road density and pastoralist residency time. Model results indicated that depredation frequency increased with the number of cattle and decreased with improved husbandry. Survey responses suggested that stronger cattle enclosures using plastic and light deterrents were husbandry techniques most supported by pastoralists. We recommend testing their efficacy and feasibility. This thesis illustrates the importance of interdisciplinary research to better inform human-carnivore coexistence grounded in the local context.

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Anthropogenic landscape change modifies the face of our planet, creating new playing fields on which wildlife communities respond to altered landscapes. Individual species react to disturbance, which trigger subsequent responses in their interactions with other species and thus propagate effects across ecological communities. In Alberta’s boreal forest, resource extraction has created a working landscape: a heterogeneous mosaic of natural and industrial features. The most pervasive industrial features are seismic lines – long trails cut for oil and gas exploration. Mammal community responses to seismic lines have contributed to population declines for the iconic woodland caribou (Rangifer tarandus caribou), thus motivating mitigation strategies in the form of habitat restoration. Although restoration is promised to recover caribou and restore landscape functionality, effective restoration should change wildlife responses to seismic lines, yet such responses are rarely evaluated. Further, interspecific interactions on the working landscape must be analyzed to understand how differential behavioural responses across species influence community dynamics.Using camera traps in northern Alberta, I investigated how large mammals respond to human landscape change at the behavioural and community levels. I first examined seismic line use by caribou, caribou predators, and caribou apparent competitors following restoration. Three years after treatment, white-tailed deer (Odocoileus virginianus) preferred unrestored seismic lines over restored lines, while wolves (Canis lupus) strongly preferred human-use lines but did not avoid restored lines. Caribou did not respond to restoration over the study period, instead preferring seismic lines in lowland habitat. I then explored interspecific interactions, assessing how distribution of wolves influenced occurrences of black bear (Ursus americanus), coyote (Canis latrans), and lynx (Lynx canadensis) at three spatiotemporal scales. All three species showed positive associations with wolves on at least one spatiotemporal level, and black bear occurrences decreased with increasing linear density while coyote and lynx occurrences increased. Overall, I demonstrated how anthropogenic landscape change – even when implemented for conservation – induces behavioural responses that can affect community interactions and thus ripple across ecological hierarchies. These results illustrate the value of multi-species monitoring to improve understanding of community interactions, especially when making single-species management decisions that ultimately influence wildlife communities as a whole.  

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