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Pest Management and the Environment 5<br />

would help realize the potential of biocontrol agents in pest management. A few studies<br />

have focused on insect population dynamics and key mortality factors under fi eld conditions<br />

(Zalucki et al., 1986; Fitt, 1989). Early-stage mortality is invariably the most severe,<br />

although its causes and extent vary greatly, and comparable data sets are too few to<br />

identify the factors responsible for population regulation across regions. Varieties with<br />

moderate levels of resistance that allow the pest densities to remain below ETLs are best<br />

suited for use in IPM in combination with natural enemies. Restless behavior and prolonged<br />

developmental period of the immature stages increases the susceptibility of the target pest<br />

to the natural enemies. However, plant morphological characteristics and secondary plant<br />

substances sometimes reduce the effectiveness of natural enemies for pest management.<br />

The use of insect-pest-resistant varieties and biological control brings together unrelated<br />

mortality factors, which reduce the pest population’s genetic response to selection pressure<br />

from plant resistance and natural enemies. Acting in concert, they provide a density-<br />

independent mortality at times of low pest density, and density-dependent mortality at<br />

times of high pest density (Bergman and Tingey, 1979).<br />

Parasitoids<br />

A diverse range of parasitoids lay their eggs on or in the body of an insect host, which is<br />

then used as a food by the developing larvae. The most important parasitoid groups are<br />

trichogrammatids, ichneumonids, braconids, chalcids, and tachinids. The trichogrammatids<br />

parasitize the eggs of several insect species, and have been used extensively in biological<br />

control. The ichneumonids and braconids prey mainly on the larvae of butterfl ies and<br />

moths. The chalcid wasps parasitize the eggs and larvae of insects. The most important<br />

and widely used parasitoids for biological control of insects are the egg parasitoids such as<br />

Trichogramma, Chelonus, and Telenomus. The larval parasitoids such as Cotesia, Encarsia,<br />

Gonatocerus, Campoletis, Bracon, Enicospilus, Palexorista, Carcelia, Sturmiopsis, etc., have also<br />

been used in several countries for biological control of insects.<br />

Cropping systems can be altered successfully to augment and enhance the effectiveness<br />

of natural enemies (Andow and Risch, 1985; Altieri, Wilson, and Schmidt, 1985). Optimal<br />

microclimatic conditions, nectar sources, and alternate hosts may exist in some cropping<br />

systems, but not others. Physicochemical characteristics of the host plants also play an<br />

important role in host specifi city of both the insect hosts and their parasitoids (Sharma,<br />

Pampapathy, and Sullivan, 2003). Host-plant-mediated differences in the activity and<br />

abundance of natural enemies have been recorded in the case of Helicoverpa armigera<br />

(Hubner) (Pawar, Bhatnagar, and Jadhav, 1986; Zalucki et al., 1986; Manjunath et al., 1989).<br />

The average rates of parasitism of the eggs of H. armigera (mainly by Trichogramma spp.)<br />

have been found to be 33% on sorghum, 15% on groundnut, and 0.3% on pigeonpea, and<br />

little or no parasitism was observed on chickpea (Pawar, Bhatnagar, and Jadhav, 1986).<br />

Manjunath et al. (1989) observed up to 98% parasitism of H. armigera eggs by Trichogramma<br />

chilonis Ishii on tomato, potato, and lucerne, but no egg parasitism was recorded on chickpea,<br />

probably because of the acid exudates secreted by the leaves. Therefore, due consideration<br />

should be given to the host plant and the species of the parasitoid involved while<br />

planning for biological control of insect pests.<br />

Predators<br />

In general, predators have received much less attention than parasites as natural control<br />

agents. They exercise greater control on pest populations in a diverse array of crops

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