Size, time, and asynchrony matter: the species-area relationship for parasites of freshwater fishes.

The tendency to attribute species-area relationships to "island biogeography" effectively bypasses the examination of specific mechanisms that act to structure parasite communities. Positive covariation between fish size and infrapopulation richness should not be examined within the typical extinction-based paradigm, but rather should be addressed from the standpoint of differences in colonization potential among individual hosts. Although most mechanisms producing the aforementioned pattern constitute some variation of passive sampling, the deterministic aspects of the accumulation of parasite individuals by fish hosts makes untenable the suggestion that infracommunities of freshwater fishes are stochastic assemblages. At the component community level, application of extinction-dependent mechanisms might be appropriate, given sufficient time for colonization, but these structuring forces likely act indirectly through their effects on the host community to increase the probability of parasite persistence. At all levels, the passive sampling hypothesis is a relevant null model. The tendency for mechanisms that produce species-area relationships to produce nested subset patterns means that for most systems, the passive sampling hypothesis can be addressed through the application of appropriate null models of nested subset structure.