Theory for Biological Control: Recent Developments

Abstract

It has been argued that ecological theory has not been useful to the practice of biological control. The purpose of this article is to show how recent theoretical advances may reverse this situation. We first discuss four issues that have arisen in the development of theory for biological control. (1) Recent work has clarified and resolved earlier disagreements concerning the effect of aggregation of the parasitoid to local host density on the stability of parasitoid—host models; this work emphasizes that such aggregation increases the ability of the parasitoid to reduce pest density. (2) There has been disagreement over the conditions under which stability, in the mathematical sense and on a local spatial scale, is an appropriate goal for classical biological control of insect pests, and whether a metapopulation may sometimes provide a more appropriate framework. We also comment on (3) relative size of refuges, which has been proposed as a unifying concept, and (4) density dependence and ratio dependence. We then discuss recent models using a stage—structured approach, particularly those that compare potential biological control agents or agents that have been differentially successful in practice. We argue that this approach has already produced valuable insights into the factors operating in several field situations. It demonstrates the importance of identifying which pest stages are most injurious to the crop, since the natural enemy that wins in competition may not be the most effective at suppressing the crucial pest stage(s): in general, the winning parasitoid reduces the density of the host stage attacked by its competitor below the level at which the latter can maintain a positive growth rate when at low density. A useful criterion is that pest equilibrium density is suppressed most by the parasitoid species that needs the fewest host individuals to allow a female parasitoid to replace herself in the next generation. Caveats are that this may apply only under equilibrium conditions and that the pest stage most suppressed is the one we wish to control. We discuss the connection between successful biological control and evolutionary considerations.