Habitat association in populations on landscapes with continuous-valued heterogeneous habitat quality

We explore a spatially implicit patch-occupancy model of a population on a landscape with continuous-valued heterogeneous habitat quality, primarily considering the case where the habitat quality of a site affects the mortality rate but not the fecundity of individuals at that site. Two analytical approaches to the model are constructed, by summing over the sites in the landscape and by integrating over the range of habitat quality. We obtain results relating the equilibrium population density and all moments of the probability distribution of the habitat quality of occupied sites, and relating the probability distributions of total habitat quality and occupied habitat quality. Special cases are considered for landscapes where habitat quality has either a uniform or a linear probability density function. For these cases, we demonstrate habitat association, where the quality of occupied sites is higher than the overall mean quality of all sites; the discrepancy between the two is reduced at larger population densities. The variance of the quality of occupied sites may be greater or less than the overall variance of habitat quality, depending on the distribution of habitat quality across the landscape. Increasing the variance of habitat quality is also shown to increase the ability of a population to persist on a landscape.