Random parasite encounters coupled with condition-linked immunity of hosts generate parasite aggregation

Parasite aggregation is viewed as a natural law in parasite-host ecology but is a paradox insofar as parasites should follow the Poisson distribution if hosts are encountered randomly. Much research has focused on whether parasite aggregation in or on hosts is explained by aggregation of infective parasite stages in the environment, or by heterogeneity within host samples in terms of host responses to infection (e.g., through representation of different age classes of hosts). In this paper, we argue that the typically aggregated distributions of parasites may be explained simply. We propose that aggregated distributions can be derived from parasites encountering hosts randomly, but subsequently by parasites being 'lost' from hosts based on condition-linked escape or immunity of hosts. Host condition should be a normally distributed trait even among otherwise homogeneous sets of hosts. Our model shows that mean host condition and variation in host condition have different effects on the different metrics of parasite aggregation. Our model further predicts that as host condition increases, parasites become more aggregated but numbers of attending parasites are reduced overall and this is important for parasite population dynamics. The effects of deviation from random encounter are discussed with respect to the relationship between host condition and final parasite numbers.