Contributing to the success of Bromus tectorum in the Intermountain West may be a mechanism called apparent competition, which occurs when one species increases the pressure of a consumer on a second species. This indirect interaction has been documented only a few times in invasive plant systems, and never in a fungal pathosystem. We examined the effects of the invasive annual Bromus tectorum and predation by the seed pathogen Pyrenophora semeniperda on seedling emergence and survival for two native grasses (Pseudoroegneria spicata and Elymus elymoides), by manipulating B. tectorum densities and P. semeniperda inoculum loads in randomized plots. Identical field studies were conducted in Skull Valley, Utah (xeric site) and Sprague, Washington (mesic site). The addition of inoculum decreased emergence of native grass seedlings at both sites and increased the amount of unemerged native seeds that were killed by P. semeniperda. Higher densities of B. tectorum decreased native grass survival at the mesic site and increased survival at the xeric site probably due to the beneficial effects of B. tectorum litter on soil moisture. At both sites, there were more B. tectorum seeds found in the seed banks in plots with high B. tectorum densities than in low-density plots. This indicates an increase in available prey for P. semeniperda. There was a much lower level of infection in B. tectorum seed bank seeds at the mesic site than at the xeric site. The high level of ungerminated native seeds killed by background levels of P. semeniperda, combined with the increase in available prey for the fungus in high-density B. tectorum plots, shows that apparent competition may play a role, along with direct competition, in the success of B. tectorum. This interaction is important to consider when dealing with control of B. tectorum.

Moose populations in southern Manitoba have declined. Although the primary cause of this decline is unclear, wolf predation is considered a main contributor because wolves have limited moose populations elsewhere. To help understand the role of wolf predation in limiting moose populations we reconstructed the proportional diet of wolves using stable isotope analysis and a Bayesian stable isotope mixing model (MixSIAR). We tested the assumptions of MixSIAR by running sensitivity analyses on trophic discrimination and prior information. We observed differences in wolf diet in areas where moose and deer coexist and are spatially separated, with changes both seasonally and annually. Our results suggest white-tailed deer were preyed on during winter, even in areas where deer are locally rare, suggesting prey catchability is more important than local prey density. Seasonal changes in prey accessibility strongly affect wolf predation rates, and manipulating alternative prey populations could mitigate predation impacts on moose. / October 2015

Human footprint - in which land is converted for human use - is a leading contributor to global habitat and biodiversity loss. The accelerated rate of human landscape change to meet our growing needs has led to the direct loss of critical habitat and shifts in species distributions, interactions, and behaviour. These altered conditions affect species’ ability to adapt to environmental stressors, while some species thrive and others decline. In North America, one ungulate has successfully invaded new habitat in conjunction with human land use – the white-tailed deer. Across the continent, the invasion of white-tailed deer has led to increased competition with other ungulate species including mule deer, moose, and woodland caribou. In regions with abundant apex predators, they have become a source of primary prey as their populations increase. The mechanisms by which deer occupy landscapes in the northern extents of their geographic range are not well studied outside of the winter months, or how deer respond behaviourally to various types of human disturbance in a predator-rich environment. To address these knowledge gaps, I examined population scale resource selection across seasons and individual movement behaviour in white-tailed deer in northeastern Alberta’s intensively developed oil and gas landscape. I used previously developed models of predator frequency to spatially extrapolate wolf and black bear occurrence across my study region as indicators of indirect predation risk. I used two approaches to habitat modeling to examine deer responses to various modes of human landscape change, including roads, seismic lines, and cut blocks in addition to predators and natural habitat. Deer were best described by cumulative effects – or the combination of all of these factors – across all seasons with proximity to linear features explaining the most variation among the parameters tested. Most prominently in winter, deer strongly selected for habitat features expected to contain abundant natural sources of forage, and linear features, despite a potential increased risk of predation by wolves – suggesting that deer make energetic trade-offs between forage availability and predation risk. At the individual level, deer significantly increased their rate of movement when occupying habitat associated with predation risk. I suggest that deer make greater energetic trade-offs during winter when mobility is limited to evade predators and energetic costs are higher. The continued use of anthropogenic features post-winter, increased rate of movement and spread of landscape occupancy by deer may allude to the importance of human disturbance in maintaining deer in northern climates. Linear corridors may be an important mechanism by which deer are able to successfully colonize new areas at the northern extents of their range. My results shed light on the drivers of deer distributions in human altered landscapes for managing populations where the invasion of deer is complicit in the decline of other ungulate species such as woodland caribou in Alberta’s boreal forest. / Graduate

<p>Plant-animal interactions are affected by both abundance and distribution of interacting species and the community context in which they occur. However, the relative importance of these factors is poorly known. I examined the effects of predator host range, environmental factors, host plant populations, plant traits and fruit abortion on the intensity of pre-dispersal seed predation in 46 host populations of the perennial herb <i>Lathyrus vernus</i>. I recorded damage by beetle pre-dispersal seed predators, mainly <i>Apion opeticum </i>and <i>Bruchus atomarius</i> with different host ranges on <i>L. vernus</i> as well as on two additional host plants. Local seed predator population size was mainly influenced by plant population size, current seed production and beetle population size in the previous year, but was not strongly affected by connectivity. The monophagous seed predator was less abundant and had lower densities than the oligophagous. Both predator species had a strong ability to track fluctuations in seed production; intensity of predation increased with relative increases in seed production. Oligophagous predation on<i> L. vernus i</i>ncreased with the abundance of alternative hosts, but presence of<i> L. vernus</i> did not affect predation on alternative hosts. Abundances and trait preferences differed among three co-occurring seed predators, but were also associated with the abundance of the other species. Overall, seed predation influenced selection on flower number. I found clear indications of seed predator offence but no obvious plant defence. The pattern of fruit abortion was associated with reduced plant fitness since the seed predator had an advanced ability to locate fruits with high probability of retention. Taken together, different factors influencing abundance of the seed predator species, different preferences, and context dependent trait selection are likely to result in complex spatio-temporal variation in overall seed losses and trait selection in the common host plant.</p>

Plant-animal interactions are affected by both abundance and distribution of interacting species and the community context in which they occur. However, the relative importance of these factors is poorly known. I examined the effects of predator host range, environmental factors, host plant populations, plant traits and fruit abortion on the intensity of pre-dispersal seed predation in 46 host populations of the perennial herb Lathyrus vernus. I recorded damage by beetle pre-dispersal seed predators, mainly Apion opeticum and Bruchus atomarius with different host ranges on L. vernus as well as on two additional host plants. Local seed predator population size was mainly influenced by plant population size, current seed production and beetle population size in the previous year, but was not strongly affected by connectivity. The monophagous seed predator was less abundant and had lower densities than the oligophagous. Both predator species had a strong ability to track fluctuations in seed production; intensity of predation increased with relative increases in seed production. Oligophagous predation on L. vernus increased with the abundance of alternative hosts, but presence of L. vernus did not affect predation on alternative hosts. Abundances and trait preferences differed among three co-occurring seed predators, but were also associated with the abundance of the other species. Overall, seed predation influenced selection on flower number. I found clear indications of seed predator offence but no obvious plant defence. The pattern of fruit abortion was associated with reduced plant fitness since the seed predator had an advanced ability to locate fruits with high probability of retention. Taken together, different factors influencing abundance of the seed predator species, different preferences, and context dependent trait selection are likely to result in complex spatio-temporal variation in overall seed losses and trait selection in the common host plant.

Cette thèse a pour objet la modélisation de systèmes multi-proies–prédateurs. Elle s’intéresse particulièrement à l’influence du comportement d’un prédateur sur les interactions indirectes entre ses proies, i.e. l’effet de l’ajout d’une proie sur la densité des autres. La théorie classique prédit l’occurrence d’effets indirects négatifs entre les proies, ou compétition apparente, résultant de l’interaction avec un prédateur commun ; des résultats plus récents identifient certains mécanismes à même d’atténuer ces effets négatifs. Nos travaux revisitent les hypothèses autour de ces mécanismes dans des systèmes composés de deux proies et de leur prédateur commun. Après avoir fixé le cadre écologique en rappelant les principaux types d’interactions directes et indirectes, nous introduisons les modèles proies-prédateurs classiques. Les travaux se concentrent ensuite sur une famille de modèles présentant de la densité-dépendance négative chez les prédateurs couplés à différentes modélisations du comportement des prédateurs lorsqu’ils sont confrontés à plusieurs types de proies. Nous montrons notamment que les interactions entre ces mécanismes peuvent inverser la compétition apparente et, contre intuitivement, accroître la densité des proies par l’intermédiaire d’un prédateur commun. Nos résultats trouvent pour partie application en lutte biologique, où il est courant de chercher à favoriser les auxiliaires en aménageant leur environnement (apport de nourriture alternative, refuge, ...). Ils suggèrent que de telles pratiques peuvent se révéler contre-productives, le contrôle des ravageurs pouvant être affaibli du fait d’une distraction de leurs prédateurs. / This thesis deals with multi-prey–predators modeling. It is particularly focused on the influence of the behavior of a predator on indirect interactions between its prey, i.e, the effect of the addition of a prey on the other prey. Classical theory predicts the occurrence of negative indirect effects between prey, or apparent competition, resulting from the interaction with a common predator. More recent results identify mechanisms that mitigate these negative effects. Our work revisits the assumptions about these mechanisms in systems composed of two prey and their common predator. After setting the ecological framework by recalling the main types of direct and indirect interactions, we introduce classical predator-prey models. The work then focuses on a family of models with predator negative density-dependence coupled with different models of predator behavior when faced with different types of prey. We show that the interactions between these mechanisms can reverse apparent competition and counter-intuitively, increase prey density through their common predator. Our results are relevant to biological control programs, where a common practice aims at fostering biological control agents by providing them with alternative food or shelters. Our theory suggests that such practices may be counter-productive, pest control being disrupted by a predator distraction effect.

We are currently faced with the global challenge of conserving biological diversity while also increasing food production to meet the demands of a growing human population. Land-use change, primarily resulting from conversion to production land, is currently the leading cause of biodiversity loss. This occurs through habitat loss, fragmentation of remaining natural habitats, and resulting edge effects. Land-sparing and land-sharing approaches have been discussed as alternative ways to engineer landscapes to mitigate biodiversity loss while meeting production objectives. However, these represent extremes on a continuum of real-world landscapes, and it will be important to understand the mechanisms by which adjacent land use affects natural remnant ecosystems in order to make local land-management decisions that achieve conservation, as well as production, objectives. This thesis investigates the impact of juxtaposing production and natural forest on the community-wide interactions between lepidopteran herbivores and their parasitoids, as mediated by parasitoid spillover between habitats. The first and overarching objective was to determine whether herbivore productivity drives asymmetrical spillover of predators and parasitoids, primarily from managed to natural habitats, and whether this spillover alters trophic interactions in the recipient habitat. The study of trophic interactions at a community level requires understanding of both direct and indirect interactions. However, community-level indirect interactions are generally difficult to predict and measure, and these have therefore remained understudied. Apparent competition is an indirect interaction mechanism thought to be very important in structuring host-parasitoid assemblages. However, this is known primarily from studies of single species pairs, and its community-wide impacts are less clear. Therefore, my second objective was to determine whether apparent competition could be predicted for all species pairs within an herbivore assemblage, based on a measure of parasitoid overlap. My third objective was to determine whether certain host or parasitoid species traits can predict the involvement of those species in apparent competition. My key findings were that there is a net spillover of generalist predators and parasitoids from plantation to native forest, and that for generalists, this depends on herbivore abundance in the plantation forest. Herbivore populations across the edge were linked by shared parasitoids in apparent competition. Consequently, an experimental reduction of herbivore density in the plantation forest changed parasitism rates in the natural forest, as predicted based on parasitoid overlap. Finally, several host and parasitoid traits were identified that can predict the degree to which host or parasitoid species will be involved in apparent competition, a finding which may have extensive application in biological control, as well as in predicting spillover edge effects. Overall, this work suggests that asymmetrical spillover between production and natural habitats occurs in relation to productivity differences, with greater movement of predators and parasitoids in the managed-to-natural forest direction. The degree to which this affected species interactions has implications for landscape design to achieve conservation objectives in production landscapes.