Interactions between sources of mortality and the evolution of parasite virulence

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Abstract
A well–known result from the theory of the evolution of virulence is the prediction that the virulence of a pathogen (i.e. the rate of parasite–induced host mortality) always evolves to higher levels when host background mortality rates increase. This prediction, however, is derived from models that assume that host mortality sources combine additively to determine the overall host mortality rate. In this paper, we suggest that such additivity is probably rare for many host–pathogen systems, and explore how the predictions for the evolution of virulence are altered when interactions between host mortality sources are incorporated into the theory. Our results indicate that if mortality–source interactions are sufficiently strong then the evolutionarily stable level of virulence can actually decrease as the background mortality rate increases. Consequently, a detailed mechanistic description of how parasites and other mortality sources combine to cause host mortality is required before reliable predictions about virulence evolution can be made. Moreover, mortality–source interactions make empirical comparisons of the virulence of different parasites a much more subtle issue.