<p>This dissertation investigates the influence of spatially explicit animal behavior active subsidy distribution patterns. Active subsidies are animal-transported consumption and resources transfers from donor to recipient ecosystems. Active subsidies influence ecosystem structure, function and services in recipient ecosystems. Even though active subsidies affect ecosystem dynamics, most ecosystem models consider the influence of spatially-explicit animal behavior on active subsidy distributions, limiting the ability to predict corresponding spatial impacts across ecosystems. Spatial subsidy research documents the need for systematic models and analyses frameworks to provide generally insights into the relationship between animal space use behavior and active subsidy patterns, and advance knowledge of corresponding ecosystem impacts for a variety of taxa and ecological scenarios.</p> <p> </p> <p>To advance spatial subsidy research, this dissertation employs a combined individual-based and movement ecology approach in abstract modeling frameworks to systematically investigate the influence of 1) animal movement behavior given mortality (chapter 2), 2) animal sociality (chapter 3) and 3) landscape heterogeneity (chapter 4) on active subsidy distribution. This dissertation shows that animal movement behavior, sociality and landscape heterogeneity influence the extent and intensity of active distribution and impacts in recipient ecosystems. Insights from this dissertation demonstrate that accounting for these factors in the development of ecosystem models will consequentially enhance their utility for predicting active subsidy spatial patterns and impacts. This dissertation advances spatial subsidy research by providing a road map for developing a comprehensive, unifying framework of the relationship between animal behavior and active subsidy distributions.</p>

Species’ ranges arise from the interplay between environmental preferences, biotic and abiotic environmental conditions, and accessibility. Understanding of – and predictive models on – species distributions often build from the assumption that these factors apply homogenously within each species, but there is growing evidence for individual variation. Here, I use movement data to investigate individual-level decisions and compromises regarding the different costs and benefits influencing individuals’ geographic locations, and the species-level spatial patterns that emerge from these. I first developed a new method that uses tracking data to quantify individual specialisation in geographic space (site fidelity) or in environmental space (environmental specialisation). Applying it to two species of albatrosses, I found evidence of site fidelity but weak environmental specialisation. My results have implications for how limited research efforts are best-targeted: if animals are generalists, effort are best spent by understanding in depth individual patterns, i.e., better to track fewer individuals for long periods of time; whereas if animals tend to be specialists, efforts should be dedicated to tracking as many individuals as possible, even if for shorter periods. I then investigated individual migratory strategies and their drivers in nine North American bird species, using ringing/recovery data. I found latitudinal redistribution of individuals within the breeding and non-breeding ranges that generally did not follow textbook patterns (‘chain migration’ or ‘leapfrog migration’). Migratory individuals tend to trade off the benefits of migration (better tracking of climatic niche; better access to resources) and its costs (increasing with migratory distance). I found that birds are more likely to remain as residents in areas with warmer winter temperatures, higher summer resource surpluses and higher human population densities (presumably because of a buffering effect of urban areas). Overall, my results highlight the importance of considering individual variation to understanding the ecological processes underpinning species’ spatial patterns.

Les patrons récursifs de déplacement, où l’individu revient à des sites déjà visités, sont très répandus. L’utilisation de la mémoire, supposée être avantageuse lorsque l’environnement est prévisible, pourrait être sous-jacente à l’émergence de ces patrons. Cependant, notre compréhension de l’interface mémoire-déplacement a jusqu'à présent été limitée par un manque de méthodes adaptées et d’investigation théorique des avantages de l’utilisation de la mémoire et des patrons qui en émergent. Au cours de cette thèse j’ai cherché à combler en partie ces manques. Je propose ici trois nouveaux cadres d'analyse des patrons récursifs de déplacement. Le premier délimite les zones les plus fréquemment revisitées par un individu, le deuxième détecte la périodicité dans les revisites de sites connus, et le troisième définit formellement et quantifie la routine de déplacement en termes de répétitivité de la séquence de déplacement, et propose un algorithme pour détecter les sous-séquences répétées. A l'aide d'un modèle individu-centré, nous montrons que l'utilisation de la mémoire dans un environnement prévisible est très avantageuse énergétiquement comparée à une stratégie de recherche sans mémoire, y compris en situation de compétition, et qu'elle mène à l'émergence de domaines vitaux stables et à la ségrégation spatiale entre individus. L'utilisation de la mémoire invalide plusieurs hypothèses très courantes faites par les études populationnelles, en menant à une forte déplétion de l’environnement, à une augmentation de la taille de la population à l’équilibre, et à une relation non linéaire entre la taille de population totale et l’intensité de compétition localement ressentie par les individus. Ainsi, ma thèse contribue à une meilleure compréhension des conséquences de la mémoire sur la valeur sélective des individus, sur les patrons de déplacement, et sur la démographie des populations. Elle propose des méthodes innovantes pour quantifier et caractériser les patrons récursifs de déplacement pouvant émerger de son utilisation. Ces méthodes devraient ouvrir de nouvelles opportunités de comparaisons entre individus de différentes populations ou espèces qui permettront le test d'hypothèses sur les pressions de sélection favorisant l'utilisation de la mémoire. / Recursive movement patterns, by which an individual returns to already visited sites, are very common. Memory use, hypothesized to be advantageous when the environment is predictable, could underlie the emergence of these patterns. However, our understanding of the memory-movement interface has been limited by two knowledge gaps. We still lack appropriate methodologies and theoretical knowledge of the advantages of memory use and of the patterns that emerge from it. During this PhD project, I aimed at filling in some of these gaps. I present here three new frameworks for the analysis of recursive movement patterns. The first one delimits the areas most frequently revisited by an individual, the second one detects periodic revisit patterns, and the third one formally defines and quantifies routine movement behaviour in terms of movement sequence repetitiveness, and presents an algorithm that detects the sub-sequences that are repeated. Using an individual-based model, we show that memory use, when the environment is predictable, is very energetically advantageous compared to foraging strategies that do not use memory, including in a situation of competition, and that it leads to the emergence of stable Home Ranges and spatial segregation between individuals. Memory use invalidates several hypotheses very commonly made in population studies, by leading to a stronger environmental depletion, to a higher equilibrium population size, and to a nonlinear relationship between the total population size and the individually-experienced intensity of competition. Therefore, my PhD thesis contributes to a better understanding of the consequences of memory use for the fitness of individuals, for movement patterns, and for population dynamics. It offers innovative methodologies that quantify and characterize recursive movement patterns that can emerge from its use. These methods should open new opportunities for the comparison of the movements of individuals from different populations and species, and thus the testing of hypotheses about the pressures that select for memory use.

Ruminant animals comprise the greatest proportion of herbivores around the world, provide essential ecosystem services and human consumable protein by consuming grass and human inedible dietary fiber. Herbivory pressure alters plant communities and species diversity, effectively making grazing animals ecosystem engineers in dynamic ecosystems. Development of advanced computer processing power coupled with biometric and ecosystem sensors may be employed in the internet of things framework to create an integrated information system designed to inform understanding of grazing system function and animal energy balance. Towards this end, I utilized Bos indicus / Bos taurus crossbred steers (n = 20) across two study sites each in consecutive calendar years and fitted them with GPS and accelerometer collar systems. Steers were grazed in improved grass pastures containing Tall Fescue (Festuca arundinacea) and Bermudagrass (Cyanodon dactylon). Forage samples were collected in a 20-m grid pattern at 35-day intervals to test nutritional composition, and NDVI maps were created using remotely sensed data collected using a UAV mounted camera system. In the first chapter, I utilize the movement ecology framework to investigate metabolic theory and animal behavior on energy budgets, then explore available technology to utilize in an integrative information system. In Chapter 2, I tested preprocessing and behavior collection methods used to train a machine learning randomforest classification model to predict animal behavior using triaxial accelerometers. Landscape functional scale and optimal sampling density is the primary focus of Chapter 3, where I explored the complex relationship between sampling regime, interpolation strategy, and landscape complexity, demonstrating that sampling density is a product of desired accuracy and landscape complexity. Finally, I focused on animal growth in Chapter 4, demonstrating the functionality of a walk-over-weigh system, and identified robust regression as the most accurate smoothing method to identify and remove spurious animal weights.

New analytical and technological tools have the potential to yield unprecedented insights into the life histories of migratory species. I used Bayesian population models and Global Positioning System-acceleration tracking devices to understand the demographic mechanism and likely drivers underpinning the Greenland White-fronted Goose (Anser albifrons flavirostris) population decline. I used a 27-year capture-mark-recapture dataset from the main wintering site for these birds (Wexford, Ireland) to construct multistate models that estimated age- and sex-specific survival and movement probabilities and found no sex-bias in emigration or ‘remigration’ rates (chapter 2). These formed the foundation for an integrated population model, which included population size and productivity data to assess source-sink dynamics through estimation of age-, site-, and year-specific survival and movement probabilities, the results of which suggest that Wexford is a large sink and that a reduction in productivity (measured as recruitment rate) is the proximate demographic mechanism behind the population decline (chapter 3). Low productivity may be due to environmental conditions on breeding areas in west Greenland, whereby birds bred at youngest ages when conditions were favourable during adulthood and the breeding year (chapter 4), and possibly mediated by links with the social system, as birds remained with parents into adulthood, forfeiting immediate reproductive success, although a cost-benefit model showed the ‘leave’ strategy was marginally favoured over the ‘stay’ strategy at all ages (chapter 5). Foraging during spring does not appear to limit breeding, as breeding and non-breeding birds did not differ in their proportion of time feeding or energy expenditure (chapter 6). Two successful breeding birds were the only tagged individuals (of 15) to even attempt to nest, suggesting low breeding propensity has contributed to low productivity. Although birds wintering in Ireland migrated further to breeding areas than those wintering in Scotland, there were no differences in feeding between groups during spring migration (chapter 7). These findings suggest that Greenland White-fronted Geese are not limited until arrival on breeding areas and the increasingly poor environmental conditions there (chapter 8). More broadly, these findings demonstrate the application of novel tools to diagnose the cause of population decline.

Given current rates of anthropogenic environmental change, combined with the increasing lethal and non-lethal mortality threat that human activities pose, there is a vital need to understand wildlife movement and behaviour in human-dominated landscapes to help inform conservation efforts and wildlife management. As long-term monitoring of wildlife populations using Global Positioning System (GPS) telemetry increases, there are new opportunities to quantify change in wildlife movement and behaviour. The objective of this PhD research is to develop novel methodological approaches for quantifying change in spatial-temporal patterns of wildlife movement and habitat by leveraging long time series of GPS telemetry and remotely sensed data. Analyses were focused on the habitat and movement of individuals in the threatened grizzly bear (Ursus arctos) population of Alberta, Canada, which occupies a human-dominated and heterogeneous landscape. Using methods in functional data analysis, a multivariate regionalization approach was developed that effectively summarizes complex spatial-temporal patterns associated with landscape disturbance, as well as recovery, which is often left unaccounted in studies quantifying patterns associated with disturbance. Next, the quasi-experimental framework afforded by a hunting moratorium was used to compare the influence of lethal (i.e., hunting) and non-lethal (i.e., anthropogenic disturbance) human-induced risk on antipredator behaviour of an apex predator, the grizzly bear. In support of the predation risk allocation hypothesis, male bears significantly decrease risky daytime behaviours by 122% during periods of high lethal human-induced risk. Rapid behavioural restoration occurred following the end of the hunt, characterized by diel bimodal movement patterns which may promote coexistence of large predators in human-dominated landscapes. A multi-scale approach using hierarchical Bayesian models, combined with post hoc trend tests and change point detection, was developed to test the influence of landscape disturbance and conditions on grizzly bear home range and movement selection over time. The results, representing the first longitudinal empirical analysis of grizzly bear habitat selection, revealed selection for habitat security at broad scales and for resource availability and habitat permeability at finer spatial scales, which has influenced potential landscape connectivity over time. Finally, combining approaches in movement ecology and conservation physiology, a body condition index was used to characterize how the physiological condition (i.e., internal state) of grizzly bears influences behavioral patterns due to costs and benefits associated with risk avoidance and resource acquisition. The results demonstrated individuals in poorer condition were more likely to engage in risky behaviour associated with anthropogenic disturbance, which highlights complex challenges for carnivore conservation and management of human-carnivore conflict. In summary, this dissertation contributes 1) a multivariate regionalization approach for quantifying spatial-temporal patterns of landscape disturbance and recovery applicable across diverse natural systems, 2) support for the growing theory that apex predators modify behavioural patterns to account for temporal overlap with lethal and non-lethal human-induced risk associated with humans, 3) an integrated approach for considering multi-scale spatial-temporal change in patterns of wildlife habitat selection and landscape connectivity associated with landscape change, 4) a cross-disciplinary framework for considering the impacts of the internal state on behavioural patterns and risk tolerance. / Graduate

Pathogens and parasites are increasingly recognized as important components within host populations, communities, and ecosystems. Parasite contributions to ecosystem function most likely manifest as density-mediated impacts of parasites on their hosts, the direct contributions of parasite biomass to a system, and via parasite-induced changes in host behavior and physiology (trait-mediated impacts). Here, a framework was constructed that can be used to conceptualize parasite contributions to ecosystem function (Chapter 1). Then the influence of parasite attack on host movement was explored to further evince the mechanistic underpinnings of trait-mediated parasite impacts (Chapter 2). Additionally, mesocosms were created across a gradient of parasitism to examine how these mechanisms are likely to unfold at larger biological scales (Chapter 3). Lastly, a series of differential equations was created to model host-parasite-ecosystem interactions and generate theoretical predictions about how and when parasites are likely to influence ecosystem processes (Chapter 4). Parasites have many characteristics of ecosystem engineers, but their role has historically been ignored. These studies begin to explore the role that parasitism may have as one of the drivers of ecosystem processes.

O lobo-guará é uma espécie de ampla distribuição na América do Sul, tendo no Brasil sua maior área de ocorrência. No entanto, as modificações das áreas naturais principalmente destinadas à agropecuária tornam a espécie vulnerável à extinção. A investigação objetivou conhecer em larga escala a área de distribuição potencial gerada por atributos ambientais favoráveis e áreas adequadas à sua ocorrência nos biomas brasileiros e investigar como a espécie responde à estrutura da paisagem, avaliando os efeitos de ambientes modificados pelo homem na sua ecologia espacial, nos padrões de atividade e na movimentação. Modelos de distribuição de espécie foram gerados pelo Maxent, utilizando uma base de pontos de localização de presença a partir de 2000 para o Cerrado (Ce), Pantanal (Pa), Mata Atlântica (MA) e Pampas (Pp) e um conjunto de onze variáveis ambientais não correlacionadas (topográficas, climáticas e paisagísticas). Para análises de ecologia espacial, das atividades e de movimentação, utilizou-se localizações de telemetria (GPS) de animais habitantes de áreas protegidas (AP), e indivíduos em paisagens modificados (AM). Análises de áreas de vida (AV) foram realizadas utilizando o estimador AKDE e associadas com classificação da paisagem local. Os modelos de distribuição do lobo-guará apresentaram uma área de distribuição potencial de 78% do total dos biomas. Apesar de possuírem grandes proporções de áreas adequadas (Ce, 90%; Pa, 93%; MA, 65% e Pp, 6%), somente um pequeno percentual (4,4% do Ce e 4,7% da MA) possui adequabilidade ambiental acima de 50%. Dos atributos que favorecem sua presença, a altitude (para todos os biomas), a precipitação (Ce e Pa), diferenças de temperatura e uso e cobertura do solo (Ma e Pp) foram os mais importantes. Em nível local, animais apresentaram média de AV de 90Km2 em AP e 41Km2 em AM, uma diferença significativa (p<0,01) com áreas diretamente proporcionais ao percentual de áreas naturais na paisagem. Ainda, apesar dos padrões regulares de atividade não mostrarem grandes mudanças, o período de repouso foi significativamente maior (p<0,01) entre os animais AM (46% do dia) que em animais AP (25% do dia). Lobos-guarás de AP e AM não apresentaram grandes diferenças no deslocamento diário com média geral de 14km caminhados por dia, com comprimentos de passos de 1Km. Diferenças no comprimento de passo foram relacionadas à composição da diversidade de contato de classes da paisagem com a proporção de ambientes naturais no passo (quanto maior as variáveis, maior o passo). Passos menores refletem menor persistência de movimento interferindo no deslocamento diário. Com os resultados desse estudo identificou-se a MA e Pa muito importantes, mas o Ce como bioma mais adequado à espécie. Foram encontrados indícios de que a estrutura de suas AV, o uso da paisagem, as atividades e movimentação são afetados pela paisagem modificada. Isso pode comprometer a viabilidade populacional, interferindo na presença em uma área e refletindo no seu potencial de distribuição. As estratégias de manejo de uso do solo, e a recuperação e conexão de áreas adequadas são urgentes e necessárias para que o lobo-guará permaneça presente e funcional nas paisagens dos biomas brasileiros. / The maned wolf has an extensive distribution range throughout South America with Brazil holding the largest portion of this area. However, the species is presently under a vulnerable status due to natural habitats alteration especially from farming and ranching. This study aimed to observe in large scale the potential distribution area indicated by favorable environmental attributes and suitable habitats to its presence within the Brazilian biomes and further to investigate how the species respond to the landscape structure, evaluating the effects of human-modified landscapes on its spatial ecology, activity patterns and movements. Species distribution models were generated using Maxent with a database of presence-only locations from 2000 though 2015, of the Cerrado (Ce), Pantanal (Pa), the Atlantic Forest (AF), and the Pampas (Pp) biomes, and eleven uncorrelated environmental variables (topographic, climatic, and landscape-based). As for the spatial ecology, activity and movement analysis, GPS-based telemetry locations were used from animals inhabiting protected (PAs) and disturbed areas (DAs). Home range (HR) analysis was performed using the AKDE estimator and then associated with the landscape-classified image. The distribution models for the maned wolf showed a potential distribution area of 78% of the total biomes range. Despite the high proportion of suitable areas (Ce, 90%; Pa, 93%; AF, 65%, and Pp, 6%), only a small percentage of the biomes (4.4% for Ce and 4.7% for AF) presented values over 50% suitable. Concerning the attributes that favor its presence, the altitude (for all the biomes), the precipitation (Ce and Pa), the temperature differences and land use (AF and Pp) were the most important. At the local scale, maned wolves showed HR sizes (average) with significant differences (p<0.01) between the 90Km2 (PA) and 41Km2 (DA) with HR size directly related to the proportion of natural areas. Although the general activity patterns were not considerably different between PA and DA, the resting periods of DA\'s animals (46% of the day) were significantly different (p<0.01) than the period of PA\'s wolves (46% of the day). Animals did not show changes on the daily movement patterns, accounting with 14km in general average and 1km of average step length (no major differences as well). The variation on the step length was related to the association of the diversity of contacts between landscape classes with the proportion of natural classes at each step (as bigger the variable values, bigger the step). As a consequence of smaller steps, constancy of movement decreases. Considering the results, the AF and Pa outstand as very important biomes, however the Ce was indicated as the most suitable biome. Furthermore, the research indicated signs that the HR structure and the landscape use, besides the activity patterns and the movement are affected by altered landscapes. This might compromise the population viability, interfering directly on its presence in an area and affecting the species distribution. Thus, a proper land use management aiming the recovering of degraded habitats is an important strategy tor the maned wolf conservation, so the species can long last survive across the Brazilian biomes.

O lobo-guará é uma espécie de ampla distribuição na América do Sul, tendo no Brasil sua maior área de ocorrência. No entanto, as modificações das áreas naturais principalmente destinadas à agropecuária tornam a espécie vulnerável à extinção. A investigação objetivou conhecer em larga escala a área de distribuição potencial gerada por atributos ambientais favoráveis e áreas adequadas à sua ocorrência nos biomas brasileiros e investigar como a espécie responde à estrutura da paisagem, avaliando os efeitos de ambientes modificados pelo homem na sua ecologia espacial, nos padrões de atividade e na movimentação. Modelos de distribuição de espécie foram gerados pelo Maxent, utilizando uma base de pontos de localização de presença a partir de 2000 para o Cerrado (Ce), Pantanal (Pa), Mata Atlântica (MA) e Pampas (Pp) e um conjunto de onze variáveis ambientais não correlacionadas (topográficas, climáticas e paisagísticas). Para análises de ecologia espacial, das atividades e de movimentação, utilizou-se localizações de telemetria (GPS) de animais habitantes de áreas protegidas (AP), e indivíduos em paisagens modificados (AM). Análises de áreas de vida (AV) foram realizadas utilizando o estimador AKDE e associadas com classificação da paisagem local. Os modelos de distribuição do lobo-guará apresentaram uma área de distribuição potencial de 78% do total dos biomas. Apesar de possuírem grandes proporções de áreas adequadas (Ce, 90%; Pa, 93%; MA, 65% e Pp, 6%), somente um pequeno percentual (4,4% do Ce e 4,7% da MA) possui adequabilidade ambiental acima de 50%. Dos atributos que favorecem sua presença, a altitude (para todos os biomas), a precipitação (Ce e Pa), diferenças de temperatura e uso e cobertura do solo (Ma e Pp) foram os mais importantes. Em nível local, animais apresentaram média de AV de 90Km2 em AP e 41Km2 em AM, uma diferença significativa (p<0,01) com áreas diretamente proporcionais ao percentual de áreas naturais na paisagem. Ainda, apesar dos padrões regulares de atividade não mostrarem grandes mudanças, o período de repouso foi significativamente maior (p<0,01) entre os animais AM (46% do dia) que em animais AP (25% do dia). Lobos-guarás de AP e AM não apresentaram grandes diferenças no deslocamento diário com média geral de 14km caminhados por dia, com comprimentos de passos de 1Km. Diferenças no comprimento de passo foram relacionadas à composição da diversidade de contato de classes da paisagem com a proporção de ambientes naturais no passo (quanto maior as variáveis, maior o passo). Passos menores refletem menor persistência de movimento interferindo no deslocamento diário. Com os resultados desse estudo identificou-se a MA e Pa muito importantes, mas o Ce como bioma mais adequado à espécie. Foram encontrados indícios de que a estrutura de suas AV, o uso da paisagem, as atividades e movimentação são afetados pela paisagem modificada. Isso pode comprometer a viabilidade populacional, interferindo na presença em uma área e refletindo no seu potencial de distribuição. As estratégias de manejo de uso do solo, e a recuperação e conexão de áreas adequadas são urgentes e necessárias para que o lobo-guará permaneça presente e funcional nas paisagens dos biomas brasileiros. / The maned wolf has an extensive distribution range throughout South America with Brazil holding the largest portion of this area. However, the species is presently under a vulnerable status due to natural habitats alteration especially from farming and ranching. This study aimed to observe in large scale the potential distribution area indicated by favorable environmental attributes and suitable habitats to its presence within the Brazilian biomes and further to investigate how the species respond to the landscape structure, evaluating the effects of human-modified landscapes on its spatial ecology, activity patterns and movements. Species distribution models were generated using Maxent with a database of presence-only locations from 2000 though 2015, of the Cerrado (Ce), Pantanal (Pa), the Atlantic Forest (AF), and the Pampas (Pp) biomes, and eleven uncorrelated environmental variables (topographic, climatic, and landscape-based). As for the spatial ecology, activity and movement analysis, GPS-based telemetry locations were used from animals inhabiting protected (PAs) and disturbed areas (DAs). Home range (HR) analysis was performed using the AKDE estimator and then associated with the landscape-classified image. The distribution models for the maned wolf showed a potential distribution area of 78% of the total biomes range. Despite the high proportion of suitable areas (Ce, 90%; Pa, 93%; AF, 65%, and Pp, 6%), only a small percentage of the biomes (4.4% for Ce and 4.7% for AF) presented values over 50% suitable. Concerning the attributes that favor its presence, the altitude (for all the biomes), the precipitation (Ce and Pa), the temperature differences and land use (AF and Pp) were the most important. At the local scale, maned wolves showed HR sizes (average) with significant differences (p<0.01) between the 90Km2 (PA) and 41Km2 (DA) with HR size directly related to the proportion of natural areas. Although the general activity patterns were not considerably different between PA and DA, the resting periods of DA\'s animals (46% of the day) were significantly different (p<0.01) than the period of PA\'s wolves (46% of the day). Animals did not show changes on the daily movement patterns, accounting with 14km in general average and 1km of average step length (no major differences as well). The variation on the step length was related to the association of the diversity of contacts between landscape classes with the proportion of natural classes at each step (as bigger the variable values, bigger the step). As a consequence of smaller steps, constancy of movement decreases. Considering the results, the AF and Pa outstand as very important biomes, however the Ce was indicated as the most suitable biome. Furthermore, the research indicated signs that the HR structure and the landscape use, besides the activity patterns and the movement are affected by altered landscapes. This might compromise the population viability, interfering directly on its presence in an area and affecting the species distribution. Thus, a proper land use management aiming the recovering of degraded habitats is an important strategy tor the maned wolf conservation, so the species can long last survive across the Brazilian biomes.