Adaptive defense of pests and switching predation can improve biological control by multiple natural enemies

One of the most important questions in biological control is whether multiple natural enemies can provide greater suppression of agricultural pests than a single best enemy. Intraguild predation (IGP) among natural enemies has often been invoked to explain failure of biological control by multiple enemies, and classical theoretical studies on IGP have supported this view. However, empirical studies are inconclusive and have yielded both positive and negative results. We extend classical models by considering anti-predator behavior of pests and diet switching of omnivorous natural enemies, and examine their effects on pest control. We assume that the pest can adaptively allocate effort toward the specific defense against each predator, and that the omnivorous natural enemy can consume disproportionately more of the relatively abundant prey (switching predation) by type III functional responses to prey items. The model predicts that adaptive defense augments pests but favors introduction of multiple natural enemies for controlling pests if IGP is weak. In contrast, switching predation does not make pest control by multiple natural enemies advantageous as in classical studies, in the absence of adaptive defense. However, switching predation reduces the necessity of defense by the pest against the omnivore and offsets the effect of adaptive defense. Thus, it makes the introduction of multiple natural enemies advantageous for pest control when the pest employs adaptive defense even if IGP is strong. These results suggest that types and combinations of behavior of prey and predators may greatly affect qualitative outcomes of biological control by multiple natural enemies.