target of the parasitoid. When the target life stage does not reveal itself by long-distance pheromonal signals, predators and parasitoids have been forced to adopt other strategies. In some systems parasitoids or predators locate herbivorous prey by exploiting plant signals induced by the herbivores (5-8). Thus, both the plants and the predators or parasitoids benefit from this interaction. In contrast to the foraging predators, some arthropods remain stationary and emit mimetic pheromone signals to attract and capture their prey (9).

During the last decade we have been investigating the chemically mediated foraging behavior of beneficial entomophagous arthropods in an effort to elucidate the factors that guide them to their hosts or prey. Our ultimate goal is to be able to manipulate and control these organisms to increase their effectiveness as biological control agents and thus reduce our dependence on pesticides for control of insect pests in agriculture. As we and our colleagues have learned more about these systems, we have found them to be quite complex in many instances. We have also found a surprising diversity of mechanisms by which these systems operate. Here we briefly survey three categories of chemically mediated predatorprey relationships which we have arbitrarily termed "eavesdropping, alarm, and deceit." Recent reviews (8-13) describe many of these systems in more detail.

EAVESDROPPING

Pheromones, by definition, are chemical signals between two members of the same species. In most instances sex pheromones are highly specific, attracting members of the same species only and not those of closely related species. However, it appears that many predatory and parasitic arthropods are able to intercept the sex pheromone signals of their prey or hosts. Bedard in 1965 (as cited in ref. 14) first reported the attraction of a parasitic wasp, the pteromalid Tomicobia tibialis Ashmead, to volatiles produced by males of the bark beetle Ips paraconfusus (Le Cônte) boring in ponderosa pine. While the identity of the volatile kairomone in this case has not been determined, it is very likely the sex or aggregating pheromone produced by the beetles. In analogous studies several bark beetle predators have been captured in traps baited with synthetic components of the pheromones of Ips and Dendroctonus species (14). Also, several hymenopterous parasites of the elm bark beetle, Scolytus multistriatus (Marsham), are attracted to components of its pheromone, multistriatin, 4-methyl-3-heptanol, and cubebene and combinations thereof (15).

Corn earworm moth, Helicoverpa zea, females emit a blend of hexadecanal and (Z)-7-, (Z)-9-, and (Z)-11-hexadecenal (16) that is a highly



The National Academies | 500 Fifth St. N.W. | Washington, D.C. 20001
Copyright © National Academy of Sciences. All rights reserved.
Terms of Use and Privacy Statement