We can compare natural communities with null models of communities to indicate how they differ from random assemblages of species (i.e., how much structure is present). However, because null models draw on observed values of species' prevalences, whatever structure already exists in natural communities affects the composition of a null model and weakens its comparative power. To address this, we developed formulae to estimate ''pre-interactive'' species prevalences permitting a more sensitive quantification of community structure. Nonetheless, if a null model deviates from the community that we base it on, it is difficult to separate the effects of heterogeneity in recruitment from competition. We have developed a method to test for each independently. Applying our analytical techniques to a well-studied guild of larval trematodes in the salt marsh snail Cerithidea californica revealed that competitive interactions among species were the most significant structuring force. Interestingly, spatial heterogeneity acted to significantly intensify species co-occurrences. This differs from previous studies, which argued that the isolating effects of spatial heterogeneity, not competition, structure these communities by reducing co-occurrences.
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