Seasonal shifts in predation habits, from a generalist in the summer to a specialist in the winter, have been documented for the great horned owl (Bulbo virginialis) in the boreal forest. This shift occurs largely due to varying prey availability. There is little study of this switching behaviour in the current literature. Since season length is predicted to change under future climate scenarios, it is important to understand resulting effects on species dynamics. Previous work has been done on a two-species seasonal model for the great horned owl and its focal prey, the snowshoe hare (Lepus americanus). In this thesis, we extend the model by adding one of the hare's most important predators, the Canadian lynx (Lynx canadensis). We study the qualitative behaviour of this model as season length changes using tools and techniques from dynamical systems. Our main approach is to determine when the lynx and the owl may invade the system at low density and ask whether mutual invasion of the predators implies stable coexistence in the three-species model. We observe that, as summer length increases, mutual invasion is less likely, and we expect to see extinction of the lynx. However, in all cases where mutual invasion was satisfied, the three species stably coexist.

The Arctic is warming faster than any other region of the globe. To conserve and manage many thousands of lakes across arctic landscapes, scientists need to understand historic and present conditions within these lakes to predict how the lakes, and the organisms that inhabit them, may respond to a changing climate. The goal of my research was to improve our understanding of what physical, chemical, and biological factors contribute to: 1) how lake food webs are assembled; and, 2) how these food webs may change in the future. First, I used long-term observations and lab experiments to determine how fish food, including zooplankton and snails, may respond to a warming climate. I then used field measurements of arctic char (Salvelinus alpinus) body characteristics, genetic samples, and fish diets to investigate if, and potentially why, populations of arctic char across a series of lakes achieve different maximum body sizes. Finally, as a method of monitoring population-level changes of fish abundance, I collected samples of arctic char DNA in lake water to test if estimated arctic char population abundances within a given lake correspond to the amount of DNA collected. Fish will require more food to eat as their metabolism increases with warming lake temperatures. Based on a thirty-year period of record, I determined zooplankton abundance increases in warmer years, indicating there is likely to be enough food for fishes in the future. Accordingly, zooplankton and snail abundance and development was also faster in warmer treatments of my lab experiments. My field observations indicated these are important prey items for arctic char. Small arctic char eat more zooplankton and large arctic char eat more snails, and these observations were consistent whether or not other predators are found in the particular lake. Similarly, my analyses did not indicate morphological or genetic differences between small and large arctic char within the same lake, suggesting arctic char size structure is determine by biological characteristics, including primary productivity and arctic char density. Indeed, estimates of arctic char population abundances across a series of lakes followed a gradient of arctic char densities, and my DNA sampling corresponded with this gradient. As there are thousands of lakes across the Arctic, my research demonstrates lake food webs, and the fishes within them, are likely to adapt to a warming climate. However, biological, chemical, and physical properties of these lakes can vary widely such that management and conservation plans may need to be developed at relatively small spatial scales across a large landscape.

京都大学 / 0048 / 新制・課程博士 / 博士(理学) / 甲第18827号 / 理博第4085号 / 新制||理||1587(附属図書館) / 31778 / 京都大学大学院理学研究科生物科学専攻 / (主査)准教授 森 哲, 教授 沼田 英治, 教授 高橋 淑子 / 学位規則第4条第1項該当 / Doctor of Science / Kyoto University / DFAM

In this thesis, we study the population dynamics of predator-prey interactions described by mathematical models with age/stage structures. We first consider fixed development times for predators and prey and develop a stage-structured predator-prey model with Holling type II functional response. The analysis shows that the threshold dynamics holds. That is, the predator-extinction equilibrium is globally stable if the net reproductive number of the predator $\mathcal{R}_0$ is less than $1$, while the predator population persists if $\mathcal{R}_0$ is greater than $1$. Numerical simulations are carried out to demonstrate and extend our theoretical results. A general maturation function for predators is then assumed, and an age-structured predator-prey model with no age structure for prey is formulated. Conditions for the existence and local stabilities of equilibria are obtained. The global stability of the predator-extinction equilibrium is proved by constructing a Lyapunov functional. Finally, we consider a special case of the maturation function discussed before. More specifically, we assume that the development times of predators follow a shifted Gamma distribution and then transfer the previous model into a system of differential-integral equations. We consider the existence and local stabilities of equilibria. Conditions for existence of Hopf bifurcation are given when the shape parameters of Gamma distributions are $1$ and $2$.

The population trajectory of woodland caribou (Rangifer tarandus caribou) in Newfoundland is currently determined by low calf survival due to high predation rates during the first 6-8 weeks after parturition. Most caribou in Newfoundland congregate and give birth in open calving grounds; consequently, in order to investigate predator-prey interactions, design research, and develop mitigation strategies, the geographic extent of the caribou calving grounds must be properly identified. We used VHF telemetry locations of caribou calves, collected from 2003-2010, to determine the spatial and temporal extent of caribou calving grounds in three study areas in Newfoundland. We put GPS collars on 47 black bears (Ursus americanus) in 3 caribou ranges where bears are having a significant impact on caribou recruitment by preying on calves during the calving season. Bear movements were greatest during the calving season, potentially increasing encounters with calves. Some bears migrated to the calving grounds just prior to caribou parturition, indicating deliberate broad-scale selection of areas of high calf density. Bears displayed interannual fidelity to calving ground usage patterns during the calving season, with some bears using the calving grounds every year, while others did not. We estimated the probability of a bear spending time in the calving grounds during the calving season as a function of the bear’s sex and mean distance to the calving grounds with logistic regression. We found that as distance increased, the odds of a bear spending time in the calving grounds decreased, and that at any given distance the odds were greater for male bears than for female bears. Our results indicate that some bears in Newfoundland are likely caribou calf predators, while others are not, and that the sex and broad-scale distribution of bears influenced the probability of a bear participating in calf predation during the calving season. The probability distribution of calf-visiting bears could be used to develop management practices to mitigate the impact of bear predation on declining caribou herds in Newfoundland.

In the Indian River Lagoon, the nudibranch Okenia zoobotryon and its egg masses are found attached to the bryozoan Zoobotryon verticillatum throughout the year. Okenia zoobotryon is thought to live, feed, and reproduce exclusively on Z. verticillatum, which would make this a very specialized predator-prey interaction. The primary goal of my study was to document certain aspects of the ecological and chemical relationships between Z. verticillatum and O. zoobotryon. Specifically, I wanted to understand the cues used by the nudibranch to locate and remain on Z. verticillatum. Population surveys on Z. verticillatum, as well as other potential hosts, were performed. From these surveys, I found a small number of O. zoobotryon adults and egg masses on the red macroalga S. filamentosa mixed with Z. verticillatum (0.01 and 0.05 g/wet-weight, respectively), none on the bryozoan Amathia distans, and none on the red macroalga Gracilaria tikvahiae. To determine if prey identification was associated with an adult cue, I ran paired-choice trials. In these, the adults did not significantly prefer Z. verticillatum. Next, laboratory culture of larvae of O. zoobotryon was performed. During larval culture some aspects of this previously undocumented life-history were recorded; adults of this species developed from planktotrophic veliger larvae that hatched out of egg masses between 4 and 6 days, the time post-hatching to settlement was 7-8 days, and metamorphosis occurred approximately 24 hours after settlement. The final question addressed in this study was; "Does O. zoobotryon feed on and take up chemicals from Z. verticillatum?" To address this question, high-performance liquid chromatography was performed on extracts of Z. verticillatum and O. zoobotryon. Both organisms were compared and found to contain similar compounds, which suggest that the nudibranch is feeding on Z. verticillatum and taking up chemicals. Concentrations of compounds in Z. verticillatum varied between populations in the northern and southern regions of the Indian River Lagoon. There are, also, differences between the chemical composition of Z. verticillatum in the IRL (northern and southern) and the California species identified by Sato and Fenical (1983).

Our goal in this thesis is to study the dynamics of the classical predator-prey model and the predator-prey model in the chemostat when a discrete delay is introduced to model the time between the capture of the prey and its conversion to biomass. In both models we use Holling type I response functions so that no oscillatory behavior is possible in the associated system when there is no delay. In both models, we prove that as the parameter modelling the delay is varied Hopf bifurcation can occur. However, we show that there seem to be differences in the possible sequences of bifurcations. Numerical simulations demonstrate that in the classical predator-prey model period doubling bifurcation can occur, possibly leading to chaos while that is not observed in the chemostat model for the parameters we use. For a delay differential equation, a prerequisite for Hopf bifurcation is the existence of a pair of pure imaginary eigenvalues for the characteristic equation associated with the linerization of the system. In this case, the characteristic equation is a transcendental equation with delay dependent coefficients. For our models, we develop two different methods to show how to find values of the bifurcation parameter at which pure imaginary eigenvalues occur. The method used for the classical predator-prey model was developed first. However, it was necessary to develop a more robust, less complicated method to analyze the predator-prey model in the chemostat with a discrete delay. The latter method was then generalized so that it could be applied to any second order transcendental equation with delay dependent coefficients. / Thesis / Doctor of Philosophy (PhD)

Accelerating rates of urbanization create novel compilations of species in urban ecosystems which experience unprecedented proximity to large human populations. Although an integral part of these urban ecosystems, mammals are difficult to study in urbanized areas. However, questions regarding urban mammalian ecology, such as the maintenance or disruption of predator-prey relationships, are vital to understanding what impacts anthropogenic factors may exert on the functions of urban ecosystems. We show that non-trapping hair equipment can effectively collect hair samples from two mammal groups – small mammals and mesopredators – in urban areas for the purpose of addressing ecological questions. We deployed non-trapping hair collection equipment at 16 different parks, preserves, and nature areas along the urbanization gradient of Philadelphia, Pennsylvania. We collected a total of 354 small mammal and 41 mesopredator hair samples. Of these samples, 58 small mammal samples and 14 mesopredator samples were of adequate weight for stable isotope analysis. We analyzed relationships between δ15N and δ13C values from each species and percent impervious surface (our proxy for urbanization) to explore potential shifts in mammal diets. Our results indicated a decoupling of predator-prey relationships between mesopredators and small mammals, particularly in urban spaces and suggested that mesopredators may be relying upon anthropogenic food subsidies. Our methodology was not only effective in addressing an ecological question, but it also provided a unique opportunity to connect with community members through QR code labels attached to our field equipment. Non-trapping hair collection methods offer a viable way to collect useful samples in urban areas while also fostering connections between community members and field research occurring in their neighborhoods. / Biology

Predicting the net effect of climate change on communities requires understanding how increasing temperatures alter interactions between predators, herbivores, and plants. Over the last several decades, warming experiments have provided important information about how species and their interactions will respond to increasing temperatures. These studies typically examine climate warming by experimentally increasing temperature at a constant level (24 hours) or asynchronously during the daytime, relative to unwarmed control treatments. However, advances in climate models now project that increases in mean global temperatures have been disproportionately driven by increasing nighttime (minimum) temperatures rather than daytime (maximum) temperatures. The timing of warming could have important ecological implications. For example, while night warming could benefit an organism by increasing temperatures towards a more thermally-optimal environment, day warming could raise temperatures beyond a thermal optimum and induce heat-stress. Consequently, mismatching the timing of warming in experiments relative to actual temperature changes could generate misleading predictions about the effects of climate warming. My dissertation has evaluated climate-warming experiments by characterizing past methods, demonstrating present methods, and providing a foundation for future studies. I conducted a meta-analysis on past terrestrial predator-prey climate warming studies that revealed experimental temperatures rarely match model projections, and the magnitude of this mismatch correlated with increased changes in measured effects. Two experiments, one focused on predator functional traits and the other trophic cascades, showed that different types of warming treatments result in different effects of climate change. The context dependency of warming effects necessitates careful consideration of experimental treatments if studies are to accurately predict the effects of climate warming. Region specific climate data are now readily available. Moving forward, ecologists can use these models to inform their warming treatments and perform experiments with the highest level of realism.

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