Many predatory arthropods eat both unparasitized herbivores and herbivores that are parasitized and contain the immature stages of endoparasitoids, a form of intraguild predation. Thus, the biological control of herbivorous arthropods can be either enhanced or disrupted by introducing a predator species to an existing host-parasitoid system. We evaluate the impact of introducing a predator, the convergent ladybird beetle, Hippodamia convergens, on the biological control of the cotton aphid, Aphis gossypii, by the parasitoid Lysiphlebus testaceipes, under field conditions. Predation on immature parasitoids by H. convergens was intense: 98-100% of aphid mummies were consumed by the end of the experiment, and H. convergens substantially reduced immature parasitoid populations. Despite the negative impact of H. convergens on aphid parasitoids, aphid population suppression was greatest in treatments containing both H. convergens and parasitoids. The parasitoid alone or in combination with H. convergens suppressed cotton aphids in a density-dependent manner and increased total plant leaf area and biomass, H. convergens did not substantially alter the percentage of aphids mummified by parasitoids and showed a partial feeding preference for unparasitized aphids over aphid mummies. We conclude that under conditions where a predator shows both a partial preference for unparasitized hosts and high levels of predation on unparasitized hosts, we may expect the predator to improve suppression of herbivores even if it produces high levels of intraguild predation. While intraguild predation is an important ecological interaction in the early-season cotton agroecosystem, it does not disrupt cotton aphid biological control.

Predators and plant resistance may act together to control herbivorous arthropod populations or antagonistically, which would reduce the control of pest populations. In a field experiment we enhanced predation by adding simulated leaf domatia to plants. Leaf domatia are small structures that often harbor predaceous arthropods that are potentially beneficial to the plant. We also manipulated host plant quality by inducing resistance with controlled, early season exposure of seedlings to spider mite herbivory.
 Our manipulations had profound consequences for the natural community of arthropods that inhabited the plants. Leaf domatia had a direct positive effect on abundances of two species of bugs and one species of thrips, all of which are largely predators of herbivores. On leaves with domatia, each of the predators was found inside the domatia two to three times more often than outside the domatia. Eggs of predaceous bugs inside leaf domatia were protected from parasitism compared to eggs outside the domatia. The positive effects of leaf domatia on predator abundances were associated with reduced populations of herbivorous spider mites, aphids, and whiteflies. Plants with experimental leaf domatia showed significantly enhanced reproductive performance.
 Induced resistance also affected the community of arthropods. Of the abundant predators, all of which also fed on the plant, only minute pirate bugs were negatively affected by induced resistance. Populations of herbivorous spider mites and whiteflies were directly and negatively affected by induction. In contrast, aphid populations were higher on plants with induced resistance compared to uninduced plants. Effects of induced resistance and domatia were additive for each of the predators and for aphids. However, spider mite and whitefly populations were not suppressed further by employing both induced resistance and domatia compared to each strategy alone. Our manipulations suggest that plant defense strategies can have positive effects on some species and negative effects on others. Negative effects of “resistance traits” on predators and positive effects on some herbivores may reduce the benefits of constitutive expression of resistance traits and may favor inducible defense strategies. Multiple plant strategies such as inducible resistance and morphological traits that aid in the recruitment of predators of herbivores may act together to maximize plant defenses, although they may also be redundant and not act additively.

Generalist predators in terrestrial arthropod communities have traditionally been viewed as predators whose dynamics are less tightly coupled to any particular prey species, but whose ecological roles are in other respects analogous to those of specialist predators. Biological-control theory for predator-prey interactions has been based upon a model of communities composed of three discrete trophic levels-plants, herbivores, and predators-in which biological control agents are top consumers and in which different species of predators interact only through competition for shared prey. Experiments employing single-plant field enclosures have suggested, however, that some generalist predators in the cotton agroecosystem function as higher-order predators, releasing populations of an herbivore, the cotton aphid Aphis gossypii, from control by another predator, the lacewing Chrysoperla carnea. Here we demonstrate through focal observations of neonate C. carnea foraging freely in the field that the high levels of mortality observed experimentally are not an artifact of cage confinement. Five generalist predators in the order Hemiptera were observed preying on neonate C. carnea. Neither cannibalism nor predation by heterospecific chrysopids was observed. The only other potential source of lacewing mortality observed was dislodgment from the plant, which occurred primarily on trichome-rich plant structures. A model of terrestrial arthropod communities incorporating higher-order predators may provide valuable insights into the regulation of herbivore populations and suggest useful avenues for biological-control research.

Generalist predators in terrestrial arthropod communities have traditionally been viewed as predators whose dynamics are less tightly coupled to any particular prey species, but whose ecological roles are in other respects analogous to those of specialist predators. Biological‐control theory for predator–prey interactions has been based upon a model of communities composed of three discrete trophic levels—plants, herbivores, and predators—in which biological control agents are top consumers and in which different species of predators interact only through competition for shared prey. Experiments employing single‐plant field enclosures have suggested, however, that some generalist predators in the cotton agroecosystem function as higher‐order predators, releasing populations of an herbivore, the cotton aphid Aphis gossypii, from control by another predator, the lacewing Chrysoperla carnea. Here we demonstrate through focal observations of neonate C. carnea foraging freely in the field that the high levels of mortality observed experimentally are not an artifact of cage confinement. Five generalist predators in the order Hemiptera were observed preying on neonate C. carnea. Neither cannibalism nor predation by heterospecific chrysopids was observed. The only other potential source of lacewing mortality observed was dislodgment from the plant, which occurred primarily on trichome‐rich plant structures. A model of terrestrial arthropod communities incorporating higher‐order predators may provide valuable insights into the regulation of herbivore populations and suggest useful avenues for biological‐control research.

Recent work in terrestrial communities has highlighted a new question: what makes a predator act as a consumer of herbivores versus acting as a consumer of other predators? Here we test three predictions from a model (Rosenheim and Corbett in Ecology 84:2538-2548) that links predator foraging behavior with predator ecology: (1) widely foraging predators have the potential to suppress populations of sedentary herbivores; (2) sit and wait predators are unlikely to suppress populations of sedentary herbivores; and (3) sit and wait predators may act as top predators, suppressing populations of widely foraging intermediate predators and thereby releasing sedentary herbivore populations from control. Manipulative field experiments conducted with the arthropod community found on papaya, Carica papaya, provided support for the first two predictions: (1) the widely foraging predatory mite Phytoseiulus macropilis strongly suppressed populations of a sedentary herbivore, the spider mite Tetranychus cinnabarinus, whereas (2) the tangle-web spider Nesticodes rufipes, a classic sit and wait predator, failed to suppress Tetranychus population growth rates. However, our experiments provided no support for the third hypothesis; the sit and wait predator Nesticodes did not disrupt the suppression of Tetranychus populations by Phytoseiulus. This contrasts with an earlier study that demonstrated that Nesticodes can disrupt control of Tetranychus generated by another widely foraging predator, Stethorus siphonulus. Behavioral observations suggested a simple explanation for the differing sensitivity of Phytoseiulus and Stethorus to Nesticodes predation. Phytoseiulus is a much smaller predator than Stethorus, has a lower rate of prey consumption, and thus has a much smaller requirement to forage across the leaf surface for prey, thereby reducing its probability of encountering Nesticodes webs. Small body size may be a general means by which widely foraging intermediate predators can ameliorate their risk of predation by sit and wait top predators. This effect may partially or fully offset the general expectation from size-structured trophic interactions that smaller predators are subject to more intense intraguild predation.