Food Webs, Risks of Alien Enemies and Reform of Biological Control

As ilwasions of alien species mount, biological control will become an increasingly important tool of conservation and agriculture. In an effort to understand indirect interactions in biological control, we review food web ecology in terms of resource competition, trophic cascades, intraguild predation, apparent competition, omnivory and a diverse set of tritrophic interactions. The most inclusive study suggests that food webs in biological control are simpler than in natural communities. Risks to non-target species created by biological control have been studied seriously for only about 20 years, and knowledge of these risks is incomplete. The greatest risks are known to be posed by the organisms with the broadest diets, such as vertebrates and the snail Euglandina rosea, which has probably caused the extinction of an entire genus of native snails in Polynesia. Some parasitoid species have been introduced that are sufficiently polyphagous to attack native insects, and cases of serious harm to non-target populations are now coming to light. However, polyphagous organisms continue to be imported for biological control. One case in point is the campaign against the Russian wheat aphid, in which over 8.5 million individual invertebrates, including more than 1 million individuals of 12 species of ladybird beetles new to North America, were released over the past 15 years, with little study of potential non-target effects, direct or indirect. Another case is the new use of the polyphagous black carp for suppression of pest snails in industrial catfish ponds. This fish poses great risks to the high native diversity of molluscs in the Mississippi drainage. We argue that risk to native flora in biological control of weeds can be judged before introduction. For the New World, the lowest non-target risk comes from stenophagous insects released against weeds with no native congeners. When weeds have native congeners, introductions of even relatively stenophagous insects have led to the use of non-target, native plants