<p>The introduction of the North American beaver (Castor canadensis) to Tierra del Fuego is a widely known example of a successful biological invasion. Beavers have impacted enormously the biodiversity of the island of Tierra del Fuego, and they are now spreading northward on the continent, prompting the governments of Argentina and Chile to seek methods to control their spread. Beavers first established in forests, where they were initially introduced, but by the 1990s they began to establish in the adjacent steppe. In this dissertation, I study the biology of invasive beavers across the two major habitat types in Patagonia and attempt to develop modeling tools that might be useful to manage their spread.</p><p>In chapter one I studied the history of the beaver introduction in Patagonia and provide evidence that the beaver introduction occurred as a single release event of 20 beavers from northern Manitoba, Canada. This not only clarifies the origin of the invasion, but also suggests that the beaver population of Patagonia descends from a smaller number of individuals than previously assumed.</p><p>In chapter two I studied the demography of invasive North American beavers in the two contrasting habitat types of the island of Tierra del Fuego, forest and steppe. Habitat differences can affect vital rates which may in turn impact the speed of the invasion, but this has been rarely addressed when managing the spread of invaders. I use repeated observations, mark-resight methods, telemetry and camera traps to estimate colony size and vital rates of beavers in the two habitats. Colony size and the number of offspring (“kits”) produced per colony per year were higher in the steppe, contrary to the belief that forest is better habitat. Here I suggest this may be the result of the longer time since invasion in the forests of Tierra del Fuego and that the forest subpopulation is showing density dependent regulation. Beaver survival was high in all age classes and was higher than survival rates recorded in North America. My work shows that beaver plasticity and predator release have likely facilitated the invasion in Patagonia. </p><p>In chapter three, I investigated the more recent invasion of beavers in an area of the Patagonian steppe. I utilized repeated high resolution satellite images to identify beaver ponds, and used them to study changes in beaver abundance and habitat use over time. The number of beaver ponds increased 85 % between 2005 and 2014. During this period, beavers changed their habitat selection pattern, presumably as a response to increased density. Beavers established on small watercourses in canyons first, but as more canyons became occupied over time, beavers moved to less preferred watercourses in plains and U-shaped valleys. Potential new beaver colonies established close to existing beaver ponds, suggesting proximity to a beaver pond is an important determinant of beaver colonization. Identifying habitat preferred by beavers in the steppe could help to increase early detection of the invader at the invasion front. This work highlights the importance of the use of high resolution remote sensing technologies to better understand and monitor biological invasions. </p><p>Finally, in chapter four, I built a spatially explicit individual-based model parameterized with data I collected in the field and use it to make management recommendations. Specifically I assessed the efficacy of a potential management strategy in which a “fire-break” (a zone beyond the current population front in which beavers are removed) perpendicular to the population front is instituted to attempt to prevent further northward spread of the beaver in continental Patagonia. I found that even a 100 km wide firebreak is insufficient to contain the spread of beavers, long dispersal events being the major cause of this failure. Further, I found that increasing the fraction of beavers culled within the firebreak does not decrease either the arrival time or the number of beavers that cross the firebreak. Counterintuitively, my model indicates that moderate levels of culling within the firebreak (rather than high) may be a more effective method to manage the invasion, likely as a result of inversely density dependent dispersal.</p> / Dissertation

Cette thèse porte sur l'analyse mathématique de modèles de compétition de plusieurs espèces microbiennes sur un seul nutriment dans un chémostat. L'objectif est de montrer la coexistence des espèces par divers mécanismes pour affirmer la biodiversité que l'on trouve dans les écosystèmes aquatiques et terrestres ainsi que dans les bioréacteurs. Nous nous somme intéressés principalement à trois mécanismes de coexistence :1- La compétition inter-spécifique entre les populations de micro-organismes et intra-spécifiques entre les individus de la même espèce.2- La floculation où l'espèce la plus compétitive inhibe sa propre croissance par la formation des flocs pour pouvoir coexister avec les autres espèces. En fait, ces bactéries en flocs consomment moins du substrat que les bactéries isolées puisqu'ils ont un moins bon accès au substrat, étant donné que cet accès au substrat est proportionnel à la surface extérieur du floc.3- La densité-dépendance dont le modèle peut être construit à partir du modèle de floculation en supposant que la dynamique de floculation est plus rapide que la croissance des espèces. Dans ce modèle densité-dépendant, le taux de croissance et le taux de prélèvement dépendent non seulement de la densité du substrat mais aussi de la densité de la biomasse.Enfin, nous avons étudié un modèle de digestion anaérobie à trois étapes avec dégradation enzymatique du substrat (matière organique) dont une partie peut être sous forme particulaire. L'analyse mathématique montre que ce modèle peut présenter la quadri-stabilité avec lessivage d'aucune, d'une ou de deux espèces selon la condition initiale. / This thesis focuses on the mathematical analysis of models of several species in competition on a single nutrient in a chemostat. The objective is to show the coexistence of microbial species by different mechanisms to affirm the biodiversity found in aquatic and terrestrial ecosystems as well as in bioreactors. We are interested mainly in three mechanisms of coexistence :1- The inter-specific competition between populations of micro-organisms and intra-specific between individuals of the same species.2- The flocculation where the species who wins the competition inhibits its growth by the formation of flocs to be able to coexist with the other species. In fact, these flocs consume less substrate than isolated bacteria since they have less access to the substrate, given that this access to the substrate is proportional to the outside surface of the floc.3- The density-dependence which the model can be construct from the flocculation model by assuming that the dynamics of flocculation is faster than the growth of the species. In this density-dependent model, the growth rate and removal rate depend not only on the density of substrate but also of the density of biomass.Finally, we studied a 3-step model of anaerobic digestion with enzymatic degradation of the substrate (organic matter) that can partly be under a solid form. The mathematical analysis shows that this model may exhibit the quadri-stability with washout of none, one or two species according to the initial condition.

Animal populations are limited by their environment and interactions with one another, the latter proportional to density. How density-dependent mechanisms regulate populations is poorly understood, particularly for migratory animals. Winter, or non-breeding, mechanisms remain particularly poorly understood for almost all migratory bird species. This dissertation tested the hypothesis that American Redstarts (Setophaga ruticilla) are regulated both within and between habitats in the winter by two mechanisms, a crowding effect and a buffer effect. As population size increases, crowding increases competition for space and resources and more individuals are relegated to lower quality habitats and forced into a transient behavioral strategy, which buffers high quality habitats from negative feedback of density. My study was carried out in Jamaica, where redstarts occupy diverse habitats. I found that population size varied among habitats, but accounting for both territorial and transient individuals was critical for accurate assessment of this variation. Ecological conditions drive differences in habitat suitability and redstart density at multiple spatial scales. Regionally, redstart survival, density, and numbers of transient individuals distinguish inland from coastal populations, consistent with differences in ecological conditions (e.g. rainfall). Locally, timing of spring departure, as well as age and sex structure distinguish neighboring habitats, consistent with increased competition for food. When population size increased over the years of this study, changes in local density were similar among habitats, and thus not the result of a buffer effect. However, increasing densities decreased food availability, which corresponded with negative feedback on departure timing and territorial behavior, i.e. more transients, in poorer habitats. This supported a crowding effect on spatial behavior (territoriality) and late-winter condition of individuals relegated to low suitability habitats. High quality inland habitats appear to buffer individuals from the effects of changing population abundance, as well as ecological conditions, evidence of a regional buffer effect. / acase@tulane.edu

<p>Tree sparrows (Passer montanus) have been studied in two areas in Sweden since 1997. At both sites, tree sparrow eggs had remarkably low hatching success. On average only 60% of the eggs hatched. Analyses have shown that this was caused by embryonic mortality, which was highly sex biased. About 70 % of the dead embryos were males, while about 65 % of all fledged nestlings were females. Impaired hatching success here related to two factors. Hatching success was lower for pairs with a male in poor body condition, and it was lower in areas with a high local population density. </p><p>A sex bias in the mortality early in life has been demonstrated in several species. Since the competitive ability of males is determined by conditions early in life, parents with poor provisioning capacity should prefer to produce female offspring in broods reared under poor conditions. The body condition of a tree sparrow during the nestling stage was well correlated to the condition as an adult, and pairs in which the male parent was in poor condition produced chicks in poor condition. Since the breeding success of a pair depended more on the condition of the male, females appear less affected by conditions early in life. Parents with poor provisioning capacity appear to bias offspring survival towards females, and a difference in the early susceptibility may be adaptive. </p><p>Changes in birth sex ratios have in some cases been suspected to result from exposure to estrogenic environmental pollutants. This was examined by exposing great- and blue tit embryos to a synthetic estrogen. Although there was a difference in the mortality rate, the difference was present also in the control groups, why this could not be attributed to estrogen exposure.</p>

Tree sparrows (Passer montanus) have been studied in two areas in Sweden since 1997. At both sites, tree sparrow eggs had remarkably low hatching success. On average only 60% of the eggs hatched. Analyses have shown that this was caused by embryonic mortality, which was highly sex biased. About 70 % of the dead embryos were males, while about 65 % of all fledged nestlings were females. Impaired hatching success here related to two factors. Hatching success was lower for pairs with a male in poor body condition, and it was lower in areas with a high local population density. A sex bias in the mortality early in life has been demonstrated in several species. Since the competitive ability of males is determined by conditions early in life, parents with poor provisioning capacity should prefer to produce female offspring in broods reared under poor conditions. The body condition of a tree sparrow during the nestling stage was well correlated to the condition as an adult, and pairs in which the male parent was in poor condition produced chicks in poor condition. Since the breeding success of a pair depended more on the condition of the male, females appear less affected by conditions early in life. Parents with poor provisioning capacity appear to bias offspring survival towards females, and a difference in the early susceptibility may be adaptive. Changes in birth sex ratios have in some cases been suspected to result from exposure to estrogenic environmental pollutants. This was examined by exposing great- and blue tit embryos to a synthetic estrogen. Although there was a difference in the mortality rate, the difference was present also in the control groups, why this could not be attributed to estrogen exposure.

Recent shifts towards ecosystem based fisheries management (EBFM) around the world have necessitated consideration of effects of fishing on a larger range of species than previously. Non-selective multispecies fisheries are particularly problematic for EBFM, as they can contribute to erosion of ecosystem structure. The trade-off between catch of productive commercial species and abundance of low-productivity species is unavoidable in most multispecies fisheries. A first step in evaluation of this trade-off is estimation of productivity of different species but this is often hampered by poor data. This thesis develops techniques for estimating productivity for data-limited species and aims to help clarify EBFM policy objectives for the fisheries of New South Wales (NSW), Australia. It begins with development of an age-structured model parameterised in terms of optimal harvest rate, UMSY. UMSY is a measure of productivity, comparable among species and easily communicated to managers. It also represents a valid threshold for prevention of overfishing. The model is used to derive UMSY for 54 Atlantic fish stocks for which recruitment parameters had previously been estimated. In most cases, UMSY was strongly limited by the age at which fish were first caught. However, for some species, UMSY was more strongly constrained by life history attributes. The model was then applied to twelve species of Australian deepwater dogshark (Order Squaliformes), known to have been severely depleted by fishing. Results showed that the range of possible values of UMSY for these species is very low indeed. These findings enabled a preliminary stock assessment for three dogsharks (Centrophorus spp.) currently being considered for threatened species listing. Preliminary results suggest they have been overfished and that overfishing continues. Finally, an Ecopath with Ecosim ecosystem model, representing the 1976 NSW continental slope, is used to illustrate trade-offs in implementation of fishing policies under alternative policy objectives. Results are compared with those of a biogeochemical ecosystem model (Atlantis) of the same system, built by scientists from CSIRO. While there were large differences in model predictions for individual species, they gave similar results when ranking alternative fishing policies, suggesting that ecosystem models may be useful for exploring broad-scale strategic management options.

Habitat selection research has focused on the role of vegetative and geologic habitat characteristics or antagonistic behavioral interactions. Conspecifics can confer information about habitat quality and provide positive density-dependent effects that may result in improved fitness, resulting in positive behavioral responses to conspecifics as a habitat selection strategy. I conducted 3 replicated, manipulative experiments to investigate use of conspecific cues in habitat selection for the golden-cheeked warbler (Dendroica chrysoparia) using simulated conspecific vocalizations during pre-settlement and post-breeding periods, across a range of woodland canopy cover. I measured territory density, pairing, and fledging success in paired treatment and control units. Territory density was >2 times higher in treatment units across the range of canopy (P = 0.02). Pairing success was positively correlated with territory density (P = 0.008). Territory density response was higher for pre-settlement than post-breeding treatment (P = 0.004). I found pre-settlement and post-breeding conspecific cues influence golden-cheeked warbler habitat selection, inducing settlement in previously unoccupied areas, and producing aggregations within areas of similar vegetative characteristics. Better understanding of social information use in habitat selection can improve our understanding of species distributions, yielding more accurate predictive distribution models; improve our ability to predict impacts of habitat changes on habitat use, survival, reproduction, and ultimately fitness; and provide a potential tool for attracting individuals to restored or managed sites.

Recent shifts towards ecosystem based fisheries management (EBFM) around the world have necessitated consideration of effects of fishing on a larger range of species than previously. Non-selective multispecies fisheries are particularly problematic for EBFM, as they can contribute to erosion of ecosystem structure. The trade-off between catch of productive commercial species and abundance of low-productivity species is unavoidable in most multispecies fisheries. A first step in evaluation of this trade-off is estimation of productivity of different species but this is often hampered by poor data. This thesis develops techniques for estimating productivity for data-limited species and aims to help clarify EBFM policy objectives for the fisheries of New South Wales (NSW), Australia. It begins with development of an age-structured model parameterised in terms of optimal harvest rate, UMSY. UMSY is a measure of productivity, comparable among species and easily communicated to managers. It also represents a valid threshold for prevention of overfishing. The model is used to derive UMSY for 54 Atlantic fish stocks for which recruitment parameters had previously been estimated. In most cases, UMSY was strongly limited by the age at which fish were first caught. However, for some species, UMSY was more strongly constrained by life history attributes. The model was then applied to twelve species of Australian deepwater dogshark (Order Squaliformes), known to have been severely depleted by fishing. Results showed that the range of possible values of UMSY for these species is very low indeed. These findings enabled a preliminary stock assessment for three dogsharks (Centrophorus spp.) currently being considered for threatened species listing. Preliminary results suggest they have been overfished and that overfishing continues. Finally, an Ecopath with Ecosim ecosystem model, representing the 1976 NSW continental slope, is used to illustrate trade-offs in implementation of fishing policies under alternative policy objectives. Results are compared with those of a biogeochemical ecosystem model (Atlantis) of the same system, built by scientists from CSIRO. While there were large differences in model predictions for individual species, they gave similar results when ranking alternative fishing policies, suggesting that ecosystem models may be useful for exploring broad-scale strategic management options.

Most animals grow substantially during their lifetime and change in competitive ability, predatory capacity and their susceptibility to predation as they grow. This thesis addresses the implications of this on regulation and dynamics within populations as well as between population interactions. In size-structured populations either reproduction or maturation may be more limiting. If juveniles are competitively superior, the competitive bottleneck will be in the adults and reproduction will be limiting. Mortality will in this case result in overcompensation in juvenile biomass through increased reproduction. Compensation in biomass was demonstrated in Daphnia pulex populations subjected to size-independent mortality, where juvenile biomass did not decrease when a substantial harvest was imposed due to increase per capita fecundity. This supported that juveniles were superior competitors and that population cycles seen in Daphnia are juvenile-driven. Compensatory responses in biomass may lead to that predators facilitate eachothers existence by feeding on a common prey, a phenomenon coined emergent facilitation. In an experimental test of the mechanism behind emergent facilitation it was demonstrated that the invertebrate predator Bythotrephes longimanus was favoured by thinning of its prey Holopedium gibberum. The thinning mimicked fish predation and targeted large individuals while Bythotrephes preferrs small prey. Size dependent predation also occurs within populations, i.e. cannibalism, were large individuals feed on smaller conspecifics. Two populations of the common guppy (Poecilia reticulata) originating from different environments were demonstrated to differ in cannibalistic degree. Cannibalism was also affected by the presence of refuges and females and juveniles from one population were better adapted to structural complexity than the other. The effects of these differences in cannibalism on population regulation and dynamics were studied in long term population experiments. Both populations were regulated by cannibalism in the absence of refuges, and displayed cannibal-driven cycles with suppression of recruitment and high population variability. The presence of refuges decreased density dependence and population variability and harvesting of large females in the absence of refuges led to population extinctions in the more cannibalistic population. The less cannibalistic population had higher population biomass and stronger density-dependence in the presence of refuges. When refuges were present, cohort competition increased and cycles with short periodicity were seen. Large individuals were not only cannibals, but could successfully prey on other species. Small and large guppies were allowed to invade resident populations of Heterandria formosa. Small invaders failed while large invaders succeeded as predation from large invaders broke up the competitive bottleneck that the resident population imposed on juveniles of the invader.

Recent shifts towards ecosystem based fisheries management (EBFM) around the world have necessitated consideration of effects of fishing on a larger range of species than previously. Non-selective multispecies fisheries are particularly problematic for EBFM, as they can contribute to erosion of ecosystem structure. The trade-off between catch of productive commercial species and abundance of low-productivity species is unavoidable in most multispecies fisheries. A first step in evaluation of this trade-off is estimation of productivity of different species but this is often hampered by poor data. This thesis develops techniques for estimating productivity for data-limited species and aims to help clarify EBFM policy objectives for the fisheries of New South Wales (NSW), Australia. It begins with development of an age-structured model parameterised in terms of optimal harvest rate, UMSY. UMSY is a measure of productivity, comparable among species and easily communicated to managers. It also represents a valid threshold for prevention of overfishing. The model is used to derive UMSY for 54 Atlantic fish stocks for which recruitment parameters had previously been estimated. In most cases, UMSY was strongly limited by the age at which fish were first caught. However, for some species, UMSY was more strongly constrained by life history attributes. The model was then applied to twelve species of Australian deepwater dogshark (Order Squaliformes), known to have been severely depleted by fishing. Results showed that the range of possible values of UMSY for these species is very low indeed. These findings enabled a preliminary stock assessment for three dogsharks (Centrophorus spp.) currently being considered for threatened species listing. Preliminary results suggest they have been overfished and that overfishing continues. Finally, an Ecopath with Ecosim ecosystem model, representing the 1976 NSW continental slope, is used to illustrate trade-offs in implementation of fishing policies under alternative policy objectives. Results are compared with those of a biogeochemical ecosystem model (Atlantis) of the same system, built by scientists from CSIRO. While there were large differences in model predictions for individual species, they gave similar results when ranking alternative fishing policies, suggesting that ecosystem models may be useful for exploring broad-scale strategic management options. / Science, Faculty of / Resources, Environment and Sustainability (IRES), Institute for / Graduate