Shared parasitoids in a metacommunity: indirect interactions inhibit herbivore membership in local communities.

The interaction between species, mediated by a shared natural enemy (i.e., apparent competition), has been the subject of much theoretical and empirical investigation. However, we lack field experiments that assess the importance of apparent competition to metacommunity structure. Here, I conducted a series of field experiments to test whether apparent competition, mediated by shared egg parasitoids (Anagrus nigriventris and A. columbi), occurs between two abundant planthopper species (Delphacodes scolochloa and Prokelisia crocea) of the North American Great Plains. The two planthoppers feed on different plant species within prairie potholes (wet depressions) and, thus, do not interact directly. At the scale of individual potholes, a five-fold pulse increase in D. scolochloa density (relative to control potholes) resulted in a steady decline in P. crocea density over two generations. As expected in cases of apparent competition, P. crocea eggs in these potholes suffered twice the level of parasitism as P. crocea eggs in control potholes. In contrast, a sixfold increase in P. crocea density had no effect on D. scolochloa density or parasitism in those potholes. The superiority of D. scolochloa over P. crocea likely can be attributed to a larger source population size, greater amount of host habitat, and/or the presence of a phenological refuge from parasitism for D. scolochloa. In another experiment, in which small populations of P. crocea were established either in close proximity to D. scolochloa or in isolation, I found that the likelihood of P. crocea persistence was 36% lower in the former than the latter populations. This difference was attributable to very high rates of parasitism of P. crocea when adjacent to D. scolochloa. These two experiments provide clear evidence that the two planthopper species engage in apparent competition and that the shared parasitoids may play a significant role in limiting membership in a local community. Based on these findings, I argue that metacommunity studies must be broadened to include higher trophic levels.