Nonrandom foraging and resource distributions affect the relationships between host density, contact rates and parasite transmission

Nonrandom foraging can cause animals to aggregate in resource dense areas, increasing host density, contact rates and pathogen transmission, but when should nonrandom foraging and resource distributions also have density-independent effects? Here, we used a factorial experiment with constant resource and host densities to quantify host contact rates across seven resource distributions. We also used an agent-based model to compare pathogen transmission when host movement was based on random foraging, optimal foraging or something between those states. Nonrandom foraging strongly depressed contact rates and transmission relative to the classic random movement assumptions used in most epidemiological models. Given nonrandom foraging in the agent-based model and experiment, contact rates and transmission increased with resource aggregation and average distance to resource patches due to increased host movement in search of resources. Overall, we describe three density-independent mechanisms by which host behaviour and resource distributions alter contact rate functions and pathogen transmission.