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Measuring Competition in a Spatially Heterogeneous Environment

This article addresses two types of spatial heterogeneity that may act to mitigate competition among species: variability in resource quality and variability in the densities of species that is not correlated with resource quality. I develop a method for quantifying these types of spatial heterogeneity both for animals that use patchily distributed resources and for annual plants. The method assesses spatial heterogeneity in terms of the expected intensity of intra- and interspecific competition experienced by individuals. A particular problem in studying the effects of spatial heterogeneity is the need to know the strength of competitive interactions across a range of resource qualities and population densities, which makes field experiments difficult. The method developed here may simplify field experiments by assessing the importance of spatial heterogeneity in a way that requires knowing only the spatial distribution of species relative to their resources, not the details of competitive interactions among species.