Excess positive associations in communities of intestinal helminths of bats: a refined null hypothesis and a test of the facilitation hypothesis.

The null hypothesis that the number of positive pairwise covariances should equal the number of negative pairwise covariances in samples from communities of randomly associated helminth species was reevaluated. The proportion of positive covariances in a sample from a community of independent species depends upon the proportion of rare species (prevalence less than 10%), the proportion of common species (prevalence greater than 90%), and the size of the sample of hosts. If rare species dominate, then there will be an excess of negative associations; if common species dominate there will be an excess of positive associations. Many helminth communities have more rare than common species, therefore samples from communities that show an equal number of positive and negative covariances have a greater number of positive associations than is expected for randomly associated species. Increased sample size will reduce the sampling bias, but at least 100 hosts are necessary and often 500-7,500 hosts are required. The excess of positive covariances between helminth species in 10 populations of bats disappeared after restricting the analyses to hosts in which both members of a species pair were present. This result suggests that excess positive associations between helminth species in bats are due to joint presences and absences in hosts rather than to interspecific facilitation. Interspecific facilitation would be supported by observed positive correlations between the intensities of individuals of the species pairs.