Influence of host-plant density and male harassment on the distribution of female Euphydryas anicia (Nymphalidae)

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Abstract
We studied behavioral mechanisms underlying the spatial distribution of Euphydryas anicia (Lepidoptera: Nymphalidae) females searching for larval host plants on a flat plain in Colorado. The rate of female movement, as represented by an empirically estimated diffusion coefficient, is affected by two factors. First, when searching in areas of high host-plant density, females tend to make sharper turns and fly for shorter distances between landings, compared to areas of low plant density. As a result, the rate of female displacement is lowered when they search in areas of high host-plant density, and thus females tend to aggregate in such areas. The second factor affecting female movement is the presence of males. Harassment by males induces females to increase the rate of their movement, and females are often chased out of host patches by males. Our results explain the observed spatial and temporal patterns of female distribution in relation to the distribution of host plants and males. In years when host plants are plentiful, harrassement by males in one host patch induces females to move to another. This pattern of female movement tends to disrupt the relationship between the spatial distributions of host plants and females, as well as females and males. In the year when hosts were concentrated in a single large patch, females could not easily avoid male harassment, and consequently we observed strong corelations of female-host distributions and male-female distributions.