Intraguild predation, invertebrate predators, and trophic cascades in lake food webs

The top-down and bottom-up properties of model food webs that include intraguild predation and self-limiting factors such as cannibalism are investigated. Intraguild predation can dampen or even reverse the top-down effects predicted by food chain theory. The degree of self-limitation among the intraguild prey is a key factor in determining the direction and strength of the top-down response. Intraguild predation and self-limiting factors can also substantially alter the bottom-up effects of enrichment. These results can help explain the disparate results of trophic cascade experiments in lakes, where cascades are usually seen when large Daphnia are the primary herbivores, but not when smaller-bodied herbivores are dominant. Top-down manipulations should cascade at least modestly to phytoplankton in those lakes whose food web can be reasonably approximated by a chain (typically, those where Daphnia is the dominant herbivore), as predicted by food chain theory. On the other hand, smaller-bodied zooplankton are often preyed upon heavily by invertebrate predators as well as by planktivorous fish, thereby introducing elements of intraguild predation into these food webs. In this case, conventional food chain theory is likely to give incorrect predictions. Very large cascade effects may be due primarily to regime shifts between intraguild predation-dominated food webs and those that more resemble food chains, rather than due to the simple food chain cascade usually considered.