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The equilibrium structure and behavior of defoliating insect systems McNamee, Peter James

Abstract

Defoliating insect systems, defined for the purposes of this thesis as being composed of insects which defoliate forest trees and the species with which they interact, such as their host trees and their natural enemy complexes, exhibit a wide variety of population behaviors. Similarly, a number of theories and models have been proposed to explain these behaviors. These theories emphasize the importance of different ecological processes, often concentrate on the defoliator and overlook the importance of other components. Also, much of the current understanding of the dynamics of these systems has come from forest pest research and management programs, tailored towards specific pest problems and often very short term in nature. This thesis develops and begins to test a general approach for the local dynamics of defoliating insect systems. This framework outlines the system components that are necessary to predict the behavior of defoliating insect systems. It includes ways in which the equilibrium structure of defoliating insect systems, defined as the number of equilibria for each system component, the population levels at which the equilibria occur, and the processes creating the equilibria, might be found. The framework also includes methods of inducing the qualitative behavior of these sys- terns, defined as the periodicity of defoliator outbreaks, the length of outbreaks, and the dynamics of other important system components between, during, and in the decline of defoliator outbreaks. The study begins with a detailed literature review of historical theories of defoliating insect system behavior and of the documented behavior patterns of these systems. Major classes of behavior are identified, as well as the various ecological processes which have been invoked to explain these behaviors. An analysis and documentation of the equilibrium structure and behavior of three defoliating defoliating insect systems, the eastern blackheaded budworm, the eastern spruce budworm, and the jack pine sawfly, are then used to develop general rules about how equilibrium structure and behavior can be explained. This analysis, coupled with the literature review, is used to develop the framework. The framework is then tested against historical defoliator population data and general syntheses of defoliating insect system research to assess its utility and predictability. The major results of the thesis are as follows. First, it appears that the structure and behavior of a defoliating insect system can be explained with five dynamic variables: the abundances of the defoliator; the foliage; the forest; the parasitoid; and the disease; and the effects of weather acting on the defoliator. Second, there appear to be 4 classes of defoliating insect system behavior. Third, the behavior that a defoliating insect system will exhibit seems to be determined by the magnitude of weather effects on defoliator survival and recruitment, the parasitoid numerical response to changing defoliator densities, the disease numerical response to changing defoliator densities, and the vulnerability of the forest to defoliation. Fourth, there seem to be four equilibrium structures the defoliator can exhibit, and one each for the parasitoid, the foliage, the forest, and the disease. Finally, the framework suggests that defoliating insect system structure and behavior can be induced with a particular, well-defined set of information. The framework is successful when applied to particular defoliating insect systems for explaining their behavior, but less successful in explaining defoliator equilibrium structure for other systems. Opportunities for more thorough testing of the framework exist if the particular types of data outlined above are gathered for defoliating insect systems. This lack of data for testing the framework make it currently difficult to clearly define those systems in which the framework is useful and those systems in which it is not. Experiments to test the framework are described and suggestions for future types of applied research on defoliating insect systems are presented.

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