Animal movement influences ecological and biogeographical dynamics, and studying it reveals helpful insights at a time when anthropogenic activities have accelerated rates of climatic and land cover change. This dissertation addresses three fundamental questions in ecology and biogeography linked to the movement and distribution of animals. First, how do animal movements affect their environments? Second, how do the effects of land use change interact with atmospheric climate change to alter species distributions? Third, how do organisms track their climatic niches through time and space? Each question is addressed with a separate study, each generating methods and results with implications for future academic work, management, and conservation.

In the first study, I tracked the daily movements of the common hippopotamus, Hippopotamus amphibius, a megaherbivore that transports nutrient-rich biomass between terrestrial and aquatic ecosystems. I developed a spatially explicit biomass transfer model that relates rates of ingestion and egestion to movement behavior states derived from the movement data. The biomass transfer model revealed the process by which H. amphibius generates patterned landscapes of nutrient removal and deposition hotspots. In addition, the model generated maps of these nutrient transfer landscapes, making it possible to explore the spatial dynamics of nutrient transfers, and showing that the amount of biomass transferred reaches levels equivalent to rates of aboveground net primary productivity. In addition to revealing the influences of H. amphibius on ecosystem ecology, this study also provided metrics of home range size, habitat use, and movement behavior useful for conservation planning. The first study provides a method for nutrient transfer mapping which could be applied to many other species, and leverages increasing quantities of high-resolution movement tracking data to map transfers of nutrients across landscapes. This can help predict the landscape-scale ecological changes resulting from the loss of animal movements that provide nutrient transfers. The approach can also be used to map other material transport dynamics, such as animal-transported seed dispersal or the movement of persistent organic pollutants.

In the second study, I used species distribution modeling to identify the interacting effects of climate and land use change on the distribution of H. amphibius. Hydrologic change is likely to result from ongoing shifts from rain-fed to irrigated agriculture across much of sub-Saharan Africa, where H. amphibius occurs. A lack of spatial data on hydrology, especially data temporally consistent with atmospheric climate datasets, has made it difficult to build species distribution models for semiaquatic species, such as H. amphibius, which are physiologically dependent on surface water. I overcame this challenge by coupling a simple hydrologic model to scenarios of land use and climate change, identifying potential effects on H. amphibius distributions. I found that increased levels of streamflow abstraction from irrigation will lead to much greater declines in H. amphibius habitat suitability than arise from scenarios of climate change alone. I also contrasted predictions of H. amphibius distributions that incorporated only atmospheric climate variables to predictions that also incorporated hydrologic variables, and found significant improvements in model performance when hydrology was incorporated. The second study provides support for using predictive variables with strong mechanistic links to the physiology or ecology of the focal species when building species distribution models. The study also outlines a way to generate surfaces of key hydrologic variables from the climate surfaces commonly used for species distribution modeling. These surfaces have the potential to greatly improve forecasts generated by other semiaquatic species distribution models. From a conservation perspective, the second study highlights the potential for substantial losses of H. amphibius habitat across Africa as a result of increases in irrigation development. Other semiaquatic species in the region, as well as those dependent on the keystone ecological role of H. amphibius and its nutrient-transporting movements, may be similarly affected.

In the third study, I explored the role of movement in shaping species distributions in variable climates. Climatic variability at multiple time scales causes suitable climatic conditions to shift across geographic space. Recent scholarship has proposed that two species traits, the ability to colonize suitable locations, referred to as dispersal, and the ability to continue to occupy an area with unsuitable conditions, referred to as persistence, facilitate niche tracking, the process by which species follow suitable conditions moving through geographic space. By developing a model that simulates niche tracking through historically observed patterns of temporal and spatial variability, I quantified how different dispersal and persistence abilities affect niche tracking potential. I found that both dispersal and persistence facilitate niche tracking, and that small increases in persistence ability result in surprisingly large increases in niche tracking potential. The third study makes two main contributions to ecological niche theory and distribution modeling. The first contribution is to extend niche theory to explicitly address niche tracking, enabling the spatially and temporally explicit mapping of niche tracking dynamics on real landscapes. The second contribution is to quantify the effects on niche tracking potential of increasing persistence and dispersal abilities across real climate surfaces. The results suggest that climate adaptation actions should focus not just on the ability of species to move in response to climate change, but also on their ability to persist through periods of unsuitable conditions.

Anthropogenic habitat fragmentation is a primary driver of species endangerment across the globe and has compounding effects on species diversity and ecosystem function. While efforts to enhance habitat connectivity are therefore essential to protecting biodiversity, a fundamental behavioral and ecological understanding of animal movement is first needed to successfully protect species movements. Understanding the role of behavior in determining animal movement patterns is essential to conservation planning, yet the extent to which an animal’s behavioral state (e.g. foraging, dispersing) influences its movements and resource selection has largely been ignored as part of conservation planning efforts. Further, because empirical studies of animal movement are most-often site- and species-specific, the processes underlying observed movement patterns are not well understood across taxa. This dissertation seeks to elucidate the processes that shape animal movements to advance the biological grounding of connectivity science and inform conservation efforts.

A systematic review of connectivity studies employing resource selection analysis examined how researchers have incorporated animal behavior into connectivity planning, and highlighted promising approaches for identifying wildlife corridors. The review revealed that most of the research done to date has superficially considered all animal location data as representative of resource selection patterns, despite recognition that an animal’s behavioral state can be an important component of space use. Those studies in the review that validated connectivity predictions with independent movement data indicate that general patterns of resource selection are not always a suitable proxy for movement preference during dispersal, and failure to recognize this distinction may have important consequences for species-specific efforts to preserve habitat connectivity.

Using high-resolution GPS and activity data from African wild dogs (Lycaon pictus), an Endangered species highly sensitive to habitat fragmentation, resource selection and responses to roads were measured across three behavioral states identified from activity data (hunting, resting, and traveling). The response of wild dogs to roads varied markedly with both the behavioral and landscape contexts in which roads were encountered, ranging between strong selection for and avoidance of roads depending on behavioral state. A comparison of these outputs to a full model that did not parse for behavior revealed that these patterns were not evident when all movement data were considered together in the full model. This study indicates that including behavioral information in resource selection models is critical to understanding wildlife responses to landscape features and suggests that successful application of resource selection analyses to conservation planning requires explicit examination of the behavioral contexts in which movement occurs.

The effects of behaviorally-mediated patterns of resource selection were then applied in a habitat connectivity modeling context. To illustrate the importance of behavioral information in connectivity assessments, behavior-specific predictions of connectivity were tested against long-distance dispersal movements of African wild dogs. Findings demonstrated that including only directed-movement behavior when measuring resource selection reveals far more accurate patterns of habitat connectivity than a model measuring resource selection independent of behavioral state. Results of this work suggest that connectivity studies that rely on resource selection analysis alone may be insufficient to target movement pathways and corridors for protection. This research highlights the value of incorporating animal behavior into connectivity planning.

To examine how basic movement processes scale up to produce emergent patterns for multiple species, movement data from over a dozen marine and terrestrial vertebrate species spanning three taxonomic classes, continents and orders of magnitude in body size were compared with computer-simulated idealized movement paths. This comparative approach revealed that similar movement patterns and properties recur in highly dissimilar ecological systems, and showed that a simple set of metrics can reliably classify broad-scale movement patterns such as migration, nomadism, and territoriality in disparate taxa. This classification system can be applied to inform predictions in multiple areas of ecological research, such as how an individual or species’ movement classification influences its response to climate change or its invasion potential in an exotic environment. In addition, this work provides researchers with a standardized set of movement metrics for expediently analyzing animal trajectories over time to detect any changes in movement pattern that may be indicative of environmental change.

Taken together, the body of work presented in this dissertation provides new approaches for researchers and practitioners to understand the ecology and conservation of animal movement, and in particular for measuring wildlife responses to widespread habitat alteration. Given limited conservation resources and rapidly changing environments, these contributions mark a key step in developing effective strategies to preserve critical wildlife movement processes.

Historical changes in climate have affected the diversity and distribution of species across the globe. Recent and rapid human-induced climate change is expected to have extreme consequences for biodiversity and ecosystem integrity. Documented species' responses to recent climate change include adaptation, distributional shifts and extinctions. There is an urgent need for scientists to improve understanding of how climate change will affect not only species distribution and abundance but also genetic diversity, which is the basis for evolutionary potential and thus critical to long-term persistence. Here I investigate the effects of climate change on the diversification, distribution and genetics of chipmunks in Sierra Nevada, California.

First, I examine the role of late Pleistocene climate fluctuations on the divergence of the endemic Alpine Chipmunk (Tamias alpinus) from its sister taxa, western populations of the Least Chipmunk (T. minimus). I use one mitochondrial gene (cytochrome b) and 14 microsatellite loci to examine the evolutionary relationship between these co-distributed species. Mitochondrial sequence data revealed that that T. alpinus and T. minimus populations share mitochondrial haplotypes with no overall geneaological separation, and that diversity at this locus is better explained by geography than by species' boundaries. In contrast, the microsatellite analysis showed that although highly differentiated populations within species exist, populations of the same species are more similar to each other than they are to members of the other species. This result suggests that the two species are distinct and there is no contemporary introgression along their parapatric boundary. Coalescent analysis of the divergence history indicated a late Pleistocene splitting time (~450ka) and subsequent, though limited, gene flow between the two lineages. The divergence of T. alpinus during this time period provides more evidence that Pleistocene glacial cycles played an important role in diversification of species in the Sierra Nevada and North America in general.

Second, I used small mammal surveys repeated over a century to assess accuracy of species distribution models in predicting known changes in chipmunk distributions, and to identify the main environmental drivers of these shifts. Historical (1900-1940) climate, vegetation and species occurrence data were used to develop single-species and multi-species multivariate adaptive regression spline (MARS) distribution models for three species of chipmunk. Models were projected onto the current (1980-2007) environmental surface and then tested against modern field resurveys of each species. I evaluated models both within and between time periods and found that even with the inclusion of biotic predictors, climate alone is the dominant predictor explaining the distribution of the study species within a time period. However, climate was not consistently an adequate predictor of the distributional change observed in all three species across time. For T. alpinus, climate alone showed the best predictive performance, strongly suggesting this species has retracted its range up in elevation due warming over the past 100 years. Modeling results showed that both climate and vegetation were factors in the collapse of T. senex populations in the study area and my modeling approach failed to predict the stability of the distribution over time observed in T. speciosus.

Third, I investigated how the climate-induced range contraction observed in T. alpinus over the past century has affected the species' overall diversity and population genetic structure in Yosemite National Park. I used one mitochondrial and seven microsatellite loci, amplified from both historical and modern specimens, to compare genetic structure of T. alpinus with that of T. speciosus, whose distribution remained stable over time. I found a decline in overall genetic diversity and a reduction of gene flow between local populations over time in T. alpinus, as expected from spatial modeling of distributions. As predicted given its relative stability through time, there were no significant genetic changes seen in T. speciosus.

Overall, my dissertation contributes to existing research on the diversification, climate change impacts, and conservation genetics of small mammals. Most importantly, this work exemplifies the value of natural history museums, and the use of historically collected data in contemporary biodiversity research.

Global declines in biological diversity are increasingly well documented and threaten the welfare and resilience of ecological and human communities. Despite international commitments to better assess and protect biodiversity, current monitoring effort is insufficient and conservation targets are not being met (e.g., Convention on Biological Diversity 2010 Target). Protected areas are a cornerstone of attempts to shield wildlife from anthropogenic impact, yet their effectiveness is uncertain. In this dissertation, I investigated the monitoring and conservation of wildlife (specifically carnivores and other larger mammals) within the context of a poorly studied savanna reserve in a tropical developing region: Mole National Park (MNP) in the West African nation of Ghana.

I first evaluated the efficacy of the park's long-term, patrol-based wildlife monitoring system through comparison with a camera-trap survey and an assessment of sampling error. I found that park patrol observations underrepresented MNP's mammal community, recording only two-thirds as many species as camera traps over a common sampling period. Agreement between methods was reasonable for larger, diurnal and social species (such as many larger ungulates and primates), but camera traps were more effective at detecting smaller, solitary and nocturnal species (particularly carnivores). Long-term patrol data were also subject to considerable sampling variation that could make interpretation of wildlife trends unreliable, and I suggest ways in which this locally based monitoring program may be improved.

Given the ecological and cultural importance of carnivore species, their propensity for human conflict, and the difficulty with which they are monitored, I assessed their status and vulnerability to extinction in MNP. Only 9 of 16 historically occurring carnivore species were detected in the camera-trap survey (covering 253 stations deployed for 5,469 trap days between October 2006 and January 2009). A hierarchical multi-species occupancy model applied to camera-trap data indicated a low overall likelihood of the presence of undetected species. Results from concurrent sign, call-in, and village surveys, as well as patrol records, provided more equivocal evidence of carnivore occurrence but supported the conclusion that many carnivores have declined and are likely functionally or fully extirpated from the park, including the top predator, lion (Panthera leo). Evidence of local human-carnivore conflict was also documented, including hunting of carnivores for traditional use and in retaliation for livestock depredation. Contrary to expectation, variation in carnivore persistence was not explained by ecological or life-history traits such as body size, home range size or fecundity, thus raising doubt as to the predictability of carnivore community disassembly.

I extended the multi-species occupancy model to test hypotheses about extrinsic influences on carnivore community dynamics in MNP. I derived spatially explicit GIS descriptors of heterogeneity in illegal hunting pressure, law enforcement patrol effort, prey biomass, and habitat productivity, and used a Bayesian modeling framework to assess support for their effects on carnivore occurrence. The framework explicitly accounted for spatial autocorrelation and variation in species- and site-specific detection probabilities. Contrary to my expectation, there was no indication of a consistent, negative effect of illegal hunting activity on spatial patterns of carnivore occurrence. By contrast, occurrence patterns of most species were positively associated with prey biomass, and several species had either positive or negative associations with riverine forest (but not with other indicators of habitat heterogeneity).

I conclude that pressure from hunting and other anthropogenic impacts remains high for West African wildlife, even within protected areas, but that human-wildlife relations are complex and their consequences inadequately predicted by simple models of extinction risk. Existing monitoring programs may generate data unsuitable for strong inference on wildlife community dynamics, and careful attention to objectives and methodology is needed. More attention to the protection and recovery of carnivore populations is also needed, as are further focused and interdisciplinary efforts to inform and improve wildlife conservation in West Africa.

Like much of the global biosphere, wildlife species have experienced rapid declines during the Anthropocene. Wildlife ecologists have responded to these crises by developing a range of technologies, techniques, and large datasets, which together have revolutionized the field, provided novel insights into the movements and behaviors of animals, and identified new risks and impacts to wildlife in a human-dominated world. While these advances have been vitally important, wildlife ecology has been slower to recognize and incorporate humans themselves into its new research domains. The chapters of this dissertation explore methods for better incorporating human behaviors, beliefs, actions, and infrastructure into the theories and approaches in wildlife ecology that have flourished in the last two decades. The research presented here demonstrates the importance of linking human beliefs and behaviors to wildlife ecology both by presenting novel findings and by showing the opportunities missed when narrow approaches are applied to complex socio-ecological problems.

In Chapter 1, I provide a general introduction on the theories underlying this research, contextualize the research questions in light of the loss and recovery of large predators, and describe the research site where I collected much of the data for this dissertation. In Chapter 2, I apply the methods of movement ecology to some of the first fine-scale telemetry data collected on rifle hunters. I draw conclusions about their individual, site-level, and regional-level hunting behaviors and discuss the broad implications of these findings for hunting management. In Chapter 3, I examine livestock-predator conflict using approaches from both ecology and the social sciences. I describe a form of selection bias that is likely widespread but unreported due to the omission of social data from ecological models of conflict, and I offer guidelines for combining and translating ecological and social research on conflict. In Chapter 4, I explore the ecological impacts of one of the most globally widespread human constructions, the fence. I show for the first time the potential extent of fencing at large scales and discuss the wide variety of ecological effects of fences for both humans and ecosystems. I further highlight biases and gaps in fence research that have thus far limited a complete understanding of the environmental effects of these features. In Chapter 5, I conclude by making recommendations regarding how research might better incorporate human perceptions, decisions, and actions into ecology.

While ecosystem and human health have been closely linked, the mechanisms through which natural and social systems interact to support human health are rarely understood. Yet the rapid transformation of Earth’s natural systems makes appreciating the drivers and ramifications of environmental change for human health particularly critical. To illuminate these links, I analyze the effects of changing resource access on household livelihoods, food security, health and nutrition. Specifically, I use the case of Lake Victoria, Kenya, to examine ways individuals and households respond to and affect fish availability. I employ both qualitative and quantitative methodologies, including a longitudinal household survey, ecological monitoring of fish catch, and qualitative in-depth interviews. My analyses examine a bi-directional relationship between fish availability and the health of people who depend on fish resources. First, I analyze how participation in fishing livelihoods affects fish consumption and food security. Next, I examine the bounds of resource availability in time and space in affecting household fish consumption. Further, I assess the impact of illness in fishers on fishing methods, and thereby fishery sustainability. Finally, I analyze how fish declines affect women’s access to fish, and, in particular, the power dynamics of transactional fish-for-sex relationships and HIV risk. Through these analyses, I aim to illuminate both the extensive effects of environmental change and mechanisms that tie the health of people and their environment.

Monitoring a species' distribution and abundance is a critical component in both applied and theoretical ecology. Wildlife managers demand effective and efficient methods for monitoring species of concern. Theoretical ecologists, as well, need accurate estimates of species' distributions and abundance, and measures of the quality of habitat that determines these parameters. Recent advances in species distribution models have suggested an inexpensive and robust way forward for estimating habitat suitability, but these models have rarely addressed species distributions at ecologically relevant spatial or temporal scales. Further, few studies have examined the relationship between habitat suitability, as estimated from species distribution models, and habitat quality, a better predictor of long-term population persistence. In this study, I (1) test current methods for monitoring giant kangaroo rat population extent and abundance; (2) create novel species distribution models that incorporate primary productivity as an ecological predictor for giant kangaroo rat distribution; and (3) test whether these models are actually related to independent measures of habitat quality. Aerial surveys are shown to be a reliable estimator for giant kangaroo rat population extent. Burrow counts may be a reliable index of long-term population size, but are inadequate to detecting short-term changes in size or growth. Distribution models that incorporate ecologically relevant variables on annual time scales are shown to better predict giant kangaroo rat distribution than static models that assume the species is at equilibrium. Finally, species distribution models perform well at measuring long-term giant kangaroo rat abundance, and temporally explicit models predict short-term abundance, but species distribution models are generally incapable of adequately distinguishing between high and low quality habitat as measured by survival or body condition.

In many of the world’s natural areas, humans now play, work, or live alongside large-bodied species of wildlife including ungulates, meso-carnivores, and even apex predators. The behavioral adjustments of these species to human activities have implications for individual fitness, population persistence, and community structure, as well as for human safety. Theory suggests that wildlife in human-dominated landscapes should modify their habitat use to avoid interactions with people, but that such avoidance may occur only in response to fine-scale spatial and temporal variation in human activity. Yet studies of the impacts of human disturbance on wildlife rarely quantify the fine-scale dynamics of human use. In this dissertation, I seek to link wildlife avoidance behavior more directly to the type, timing, intensity, and spatial distribution of human activity, thereby informing efforts to preserve relatively undisturbed spaces for large mammals in natural ecosystems regularly used by people.

I first consider spatiotemporal use of the landscape by grizzly bears (Ursos arctos) in areas of high recreation in Kananaskis Country, Alberta, Canada. For each day of the active bear season, I quantified numbers of people and vehicles using all trails, roads, and facilities located within the home ranges of GPS-collared bears. I estimated human disturbance at bear GPS positions as a function of both distance to human-use features and the average daily use on those features. Analyses revealed that when bears were in habitats adjacent to recreation infrastructure, they modified their behavior in response to daily, weekly, and seasonal fluctuations in human activity, avoiding the times and places of highest recreation. Bears responded to recreation patterns that were spatially and temporally consistent, highlighting the need to preserve predictable patterns of human use in the study area. This research demonstrates the value of quantifying fine-scale dynamics of human activity and focusing on areas of high overlap between wildlife and people to elucidate avoidance behavior of wild animals in human-dominated landscapes.

Further, I evaluate the effect of incorporating fine-scale estimates of recreation intensity on the predictive accuracy of resource selection function models developed to quantify habitat use of GPS-collared grizzly bears. Models that included static proxies for human disturbance were compared against those that estimated daily numbers of recreationists and vehicles within bear home ranges. When bears were in close proximity to trails, roads, and facilities, top models were those that rigorously quantified human use, indicating that bear habitat selection was significantly influenced by the fine-scale dynamics of recreation activity. My results suggest that when spatial overlap between human activity and wildlife is high, static representations of human disturbance may be less effective for describing wildlife behavior.

Last, I expand my focus to multiple species. Spatiotemporal patterns of occurrence of large mammals, recreationists, and domestic dogs were assessed using camera traps deployed within critical wildlife habitat bordering the town of Canmore, Alberta. Recreation was categorized by type of user, and daily numbers of recreationists and domestic dogs were quantified over a twenty-month period. Coyotes (Canis latrans) demonstrated the clearest temporal shifts in response to recreation intensity, and hikers and off-leash dogs spatially displaced several species. Results also suggested that study species responded most to past rather than current levels of human activity. My findings underscore the importance of spatial scale in recreation impact studies and indicate that recreation can have measurable effects on fine-scale habitat use and diel activity of large mammals.

The research presented in this dissertation adds to the growing body of literature on wildlife behavioral responses to human disturbance in general and outdoor recreation in particular. Human activity is increasing in most natural ecosystems, and my work suggests timely new approaches for quantifying human use and measuring its impacts on wildlife behavior in landscapes where presence of people is widespread.

The global expansion of human activity has had profound consequences for wildlife. Research has documented the effects of widespread habitat destruction and defaunation on species and ecosystems, but the more subtle pathways through which humans alter the natural world have largely escaped quantification. Much like apex predators, humans can instill strong fear in wild animals, which may adjust their activity to avoid contact with humans. In this dissertation, my collaborators and I examine pathways through which human disturbance, predation risk, and environmental heterogeneity influence animal behavior and distribution. We review the literature and synthesize theory to develop a novel framework for studying landscapes of fear, and we apply this framework in a global meta-analysis and field studies from California and Mozambique to understand how large mammals perceive and respond to spatial and temporal patterns of risk from humans and carnivores. We consider links between risk and response in complex systems with multiple predators or multiple prey species, and we explore ecology of fear dynamics in the context of seasonality, human disturbance, and restoration. Together, this work integrates disciplines of behavioral ecology, community ecology, and landscape ecology, applying predator-prey theory to understand the role of humans in ecological communities. By elucidating behavioral pathways linking human disturbance to wildlife community dynamics, this research contributes to our understanding of wildlife ecology in human-dominated landscapes and highlights mechanisms for human-wildlife coexistence.

In a telecoupled world, people and ecosystems are intricately linked across vast spatial scales. Consumption patterns, disease, and other factors on one side of the world often reverberate powerfully to shape landscapes, socio-cultural practices, and wildlife distributions in farremoved locales. These new realities require sustainability to be fundamentally achieved at global scales in order to have lasting sustainability for people and wildlife at local and regional systems. To consider these themes, my collaborators and I study a socio-ecological system of Indigenous hunting communities and bearded pigs, a cultural keystone species, in Malaysian Borneo. In Chapter 1, we build an intellectual framework that considers the ways that telecoupling unfolds in Sabah and Sarawak, driving oil palm expansion, deforestation, and socialchange. In Chapter 2, we carry out single-season, single-species occupancy models for bearded pigs that show the association of both socio-cultural factors (e.g. ethnicity and hunting accessibility) as well as environmental factors (e.g. protected area status and elevation) on bearded pig distributions across Malaysian Borneo. In Chapter 3, we document ways that oil palm plantations and urbanization have changed bearded pig hunting practices among Indigenous Kadazandusun-Murut (KDM) hunters in Sandakan District, Sabah. We also describe the ways that some hunting motivations, such as meat provision and gift-giving, have endured despite widespread changes in land-use. In Chapter 4, we investigate intergenerational hunting knowledge transfer within KDM hunting communities, also in Sandakan District, Sabah. Almost two-thirds of respondents had not, or were not intending, to pass on their hunting knowledge to their children. Moreover, many respondents reported low hunting interest among the younger generation, suggesting diminished importance of hunting practices in the future among KDM communities in Sabah. Together, our findings highlight the salience of social practices and Indigenous knowledge for bearded pig distributions in Malaysian Borneo. Our results also raise important questions about the nature of conservation values and human-wildlife interactions in a world in which hunting practices, and connection to nature more broadly, are in decline. Ancient socio-ecological links—including hunting, recreation, and other social practices—highlight profound connection points for long-term sustainability of human cultures, biophysical landscapes, and wildlife communities.