The mountain pine beetle (MPB, Dendroctonus ponderosae), a tree-killing bark beetle, has historically been part of the normal disturbance regime in lodgepole pine (Pinus contorta) forests. In recent years, warm winters and summers have allowed MPB populations to achieve synchronous emergence and successful attacks, resulting in widespread population outbreaks and resultant tree mortality across western North America. We develop an age-structured forest demographic model that incorporates temperature-dependent MPB infestations: the Susceptible-Infested-Juvenile (SIJ) model. Stability of fixed points is analyzed as a function of population growth rates, and indicates the existence of periodic outbreaks that intensify as growth rates increase. We devise analytical methods to predict outbreak severity and duration as well as outbreak return time. To assess the vulnerability of natural resources to climate change, we develop a thermally-driven mechanistic model to predict MPB population growth rates using a distributional model of beetle phenology in conjunction with criteria for successful tree colonization. The model uses projected daily minimum and maximum temperatures for the years 2025 to 2085 generated by three separate global climate models. Growth rates are calculated each year for an area defined by latitude range 42° N to 49° N and longitude range 108° W to 117° W on a Cartesian grid of approximately 4km resolution. Using these growth rates, we analyze how the optimal thermal window for beetle development is changing with respect to elevation as a result of climate change induced warming. We also use our combined model to evaluate if thermal regimes exist that would promote life cycle bivoltinism and discuss how yearly growth rates would change as a result. Outbreaks of MPB are largely driven by host tree stand demographics and spatial effects of beetle dispersal. We augment the SIJ model to account for the spatial effects of MPB dispersal throughout a forest landscape by coupling it with a Gaussian redistribution kernel. The new model generates a train of sustained solitary waves of infestation that move through a forest with constant speed. We convert the resulting integrodifference equation into a partial differential equation and search for travelling wave solutions. The resulting differential equation provides predictions of the shape of an outbreak wave profile and of peak infestation as functions of wave speed, which can be calculated analytically. These results culminate in the derivation of an explicit formula for predicting the severity of an outbreak based on the net reproductive rate of MPB and host searching efficiency.

This dissertation is concerned with effects dispersal and forest structure on forest insect population dynamics, and with identifying generating processes by comparing observed patterns to model predictions. In chapter 2, we investigated effects of changing forest landscape patterns on integro-difference models of host-parasitoid population dynamics. We demonstrated that removing habitat can increase herbivore density when herbivores don't disperse far, and parasitoids disperse further, due to differences in dispersal success between trophic levels. This is a novel potential explanation for why forest fragmentation increases the duration of forest tent caterpillar outbreaks. To better understand spatial model behaviour, we proposed a new local variation of the dispersal success approximation. The approximation successfully predicts effects of habitat loss and fragmentation on realistically complex landscapes, except when outbreak cycle amplitude is very large. Local dispersal success is useful in part because parameters can be estimated from widely available habitat data. In chapter 3, we investigated how well a discretized integro-difference model of mountain pine beetle population dynamics predicted the occurrence of new infestations in British Columbia. We found that a model with a large dispersal kernel, and high emigration from new, low severity infestations yielded the best predictions. However, we do not believe this to be convincing evidence that many beetles disperse from new, low severity infestations. Rather, we argued that differences in habitat quality, detection errors, and Moran effects can all confound dispersal patterns, making it difficult to infer dispersal parameters from observed infestation patterns. Nonetheless, predicting infestation risk is useful, and large kernels improve predictions. In chapter 4, we used generalized linear mixed models to characterize spatial and temporal variation in the propensity of jack pine trees to produce pollen cones, and account for confounding effects on the relationship between pollen cone production and previous defoliation by jack pine budworm. We found effects of stand age, and synchronous variation in pollen cone production among years. Accounting for background patterns in pollen cone production clarified that pollen cone production declines in with previous defoliation, as expected.

This dissertation is concerned with effects dispersal and forest structure on forest insect population dynamics, and with identifying generating processes by comparing observed patterns to model predictions. In chapter 2, we investigated effects of changing forest landscape patterns on integro-difference models of host-parasitoid population dynamics. We demonstrated that removing habitat can increase herbivore density when herbivores don't disperse far, and parasitoids disperse further, due to differences in dispersal success between trophic levels. This is a novel potential explanation for why forest fragmentation increases the duration of forest tent caterpillar outbreaks. To better understand spatial model behaviour, we proposed a new local variation of the dispersal success approximation. The approximation successfully predicts effects of habitat loss and fragmentation on realistically complex landscapes, except when outbreak cycle amplitude is very large. Local dispersal success is useful in part because parameters can be estimated from widely available habitat data. In chapter 3, we investigated how well a discretized integro-difference model of mountain pine beetle population dynamics predicted the occurrence of new infestations in British Columbia. We found that a model with a large dispersal kernel, and high emigration from new, low severity infestations yielded the best predictions. However, we do not believe this to be convincing evidence that many beetles disperse from new, low severity infestations. Rather, we argued that differences in habitat quality, detection errors, and Moran effects can all confound dispersal patterns, making it difficult to infer dispersal parameters from observed infestation patterns. Nonetheless, predicting infestation risk is useful, and large kernels improve predictions. In chapter 4, we used generalized linear mixed models to characterize spatial and temporal variation in the propensity of jack pine trees to produce pollen cones, and account for confounding effects on the relationship between pollen cone production and previous defoliation by jack pine budworm. We found effects of stand age, and synchronous variation in pollen cone production among years. Accounting for background patterns in pollen cone production clarified that pollen cone production declines in with previous defoliation, as expected.

The western spruce budworm (Choristoneura occidentalis Freeman) is the most widespread and destructive defoliator of Douglas-fir (Pseudotsuga menziesii (Mirb.) Franco) forests in British Columbia. Over the past two decades, western spruce budworm outbreaks have been sustained and widespread in the interior of British Columbia, leaving the forest industry and many forest-dependent communities increasingly vulnerable to the economic consequences of these outbreaks. While a great deal is known about the impact of western spruce budworm outbreaks on tree growth and form, substantial knowledge gaps remain as to the historic variability of western spruce budworm outbreaks and the consequences of defoliation on fundamental characteristics such as wood structure. This research focused on describing historic and contemporary western spruce budworm outbreaks across multiple spatial and temporal scales in south-central British Columbia using dendrochronology and wood anatomy techniques. Outbreak histories over the past 435 years were reconstructed using a network of tree-ring chronologies from central British Columbia, revealing that 12 western spruce budworm outbreaks have occurred since the early 1600s, with a mean return interval of 30 years. Further, the research illustrates that outbreaks observed over the last 40 years are not unprecedented, which does not support the perception that western spruce budworm is moving northward into central British Columbia. To evaluate the effects of a single western spruce budworm outbreak on the anatomical characteristics of Douglas-fir stemwood, tree ring data was collected from permanent sample plots that sustained both periodic and chronic western spruce budworm feeding. In mature even-aged stands of Douglas-fir, a documented outbreak occurred from 1976 to 1980 in the coastal transition zone of southern British Columbia. Based on microscopic wood anatomical measurements it was shown that the tree rings formed during this outbreak had significantly lower percentages of latewood, reduced mean cell wall thickness and smaller radial cell diameters relative to wood formed during periods without budworm feeding. Western spruce budworm defoliation temporarily modified cellular characteristics, which has implications for wood quality. In uneven-aged stands of mature Douglas-fir, located in the xeric southern interior of British Columbia, there has been a sustained western spruce budworm outbreak since 1997. Tree rings formed during this outbreak had progressively larger earlywood lumen area and radial cell diameter, reduced latewood cell wall thickness, latewood radial cell diameters, and lower percent latewood. Mixed-effects models revealed that climatic variables, defoliation severity, defoliation duration, and in limited cases canopy class were the best predictors of xylem features. The severity and duration of western spruce budworm defoliation, as well as site factors that influence moisture conditions effect the degree and direction of anatomical changes in the stemwood of Douglas-fir. This research fills a number of knowledge gaps by providing insights into the temporal and spatial dynamics of western spruce budworm outbreaks in central British Columbia over multiple centuries, and the plasticity of anatomical features in the stemwood of Douglas-fir during discrete western spruce budworm outbreaks. These research findings suggest that Douglas-fir forests are resilient to western spruce budworm outbreaks over space and time. / Graduate