Plants infested with a single herbivore species can attract natural enemies through the emission of herbivoreinduced plant volatiles (HIPVs). However, under natural conditions plants are often attacked by more than one herbivore species. We investigated the olfactory response of a generalist predators Macrolophus caliginosus to pepper infested with two-spotted spider mites, Tetranychus urticae, or green peach aphid, Myzus persicae, vs. plants infested with both herbivore species in a Y-tube olfactometer set up. In addition, the constituents of volatile blends from plants exposed to multiple or single herbivory were identified by gas chromatography-mass spectrometry (GC-MS). The mirid bugs showed a stronger response to volatiles emitted from plants simultaneously infested with spider mites and aphids than to those emitted from plants infested by just one herbivore, irrespective of the species. Combined with results from previous studies under similar conditions we infer that this was a reaction to herbivore induced plant volatiles. The GC-MS analysis showed that single herbivory induced the release of 22 additional compounds as compared with the volatiles emitted from clean plants. Quantitative analyses revealed that the amount of volatile blends emitted from pepper infested by both herbivores was significantly higher than that from pepper infested by a single herbivore. Moreover, two unique substances were tentatively identified (with a probability of 94% and 91%, respectively) in volatiles emitted by multiple herbivory damaged plants: a-zingiberene and dodecyl acetate.

The performance of herbivores, natural enemies and their interactions may be affected directly or indirectly by host plant traits, e.g. the physical plant characteristics may influence the search pattern and the functional response of predators. We studied the functional response of adult females of the predatory mite Neoseiulus cucumeris to first instar larvae of Thrips tabaci on three host plants (sweet pepper, eggplant and cucumber). The 24-h leaf disc experiments conducted at 25 ± 1°C, 60 ± 10% relative humidity and 16 : 8 h (light : dark) showed that N. cucumeris exhibited a type II functional response on all host plants. The following search rates and handling times were estimated from fitting the data to the disc equation 0.043/h and 1.798 h (cucumber); 0.048/h and 1.030 h (sweet pepper) and 0.0441/h and 2.294 h (eggplant) giving an estimated maximum predation of 13.35, 23.31 and 10.46 larvae per day respectively. The data from sweet pepper could also be described by the random predator equation (a¢: 0.051/h; T h : 0.472 h). The host plant species interacted significantly with prey density on the functional response of N. cucumeris with the relative differences in the number of thrips eaten on each host plant increasing with density. It is suggested that it is mainly the difference in trichome density between the three host plants that is responsible for the observed differences in the functional response of N. cucumeris. These results emphasize the importance of the host plant characteristics on the performance of natural enemies and for optimizing their use in biological control of pests.

Trichogramma species (Hymenoptera: Trichogrammatidae) are haplo-diploid egg parasitoids that are frequently used as biological control agents against lepidopteran pests. These wasps display two reproductive modes, including arrhenotoky (bisexuality) and thelytoky (unisexuality). Thelytokous forms are often associated with the presence of endosymbiotic Wolbachia bacteria. The use of thelytokous wasps has long been considered as a way to enhance the efficacy of biological control. The present study investigates the potential of a thelytokous Wolbachia-infected and an arrhenotokous uninfected Trichogramma brassicae Bezdenko strain as inundative biocontrol agents by evaluating their functional response towards different egg densities of the factitious host, the Angoumois grain moth, Sitotroga cerealella (Olivier) (Lepidoptera: Gelechiidae). The results revealed a type II functional response for both strains in which parasitism efficiency decreases with host egg density because of an increasing host handling time. A model with an indicator variable was used to compare the parameters of Holling's disc equation in different data sets. It was demonstrated that the two strains did not differ in host attack rate. However, the Wolbachia-infected strain did have an increased host handling time when compared to the bisexual strain. Some applied aspects of the findings are discussed.

Various physiological effects of Wolbachia infection have been reported in invertebrates, but the impact of this infection on behavior and the consequences of these behavioral modifications on fitness have rarely been studied. Here, we investigate the effect of Wolbachia infection on the estimation of host nutritive resource quality in a parasitoid wasp. We compare decision-making in uninfected and Wolbachia-infected strains of Trichogramma brassicae Bezdenko (Hymenoptera: Trichogrammatidae) on patches containing either fresh or old host eggs. For both strains, fresh eggs were better hosts than older eggs, but the difference was smaller for the infected strain than for the uninfected strain. Oviposition behavior of uninfected wasps followed the predictions of optimal foraging theory. They behaved differently toward high-vs. low-quality hosts, with more hosts visited and more ovipositions, fewer high-quality hosts used for feeding or superparasitism, and a sex ratio that was more biased toward females in patches containing high-quality hosts than in patches containing low-quality ones. Uninfected wasps also displayed shorter acceptance and rejection times in high-quality hosts than in hosts of lower quality. In contrast, infected wasps were less efficient in evaluating the nutritive quality of the host (fresh vs. old eggs) and had a reduced ability to discriminate between unparasitized and parasitized hosts. Furthermore, they needed more energy and therefore engaged in host feeding more often. This study highlights possible decision-making manipulation by Wolbachia, and we discuss its consequences for Wolbachia fitness.

Plant resistance to insect pests based on recombinant proteinase inhibitors (Pis) could interfere with natural enemies of target pests, as their own proteolytic systems may also be sensitive to large spectrum PIs. Oryzacystatin I (OCI) is a potential insect pest resistance factor currently engineered into a variety of crop plants, including potato Solanum tuberosum. Potential for OCI interfering with female reproduction in Perillus bioculatus, a stinkbug predator of Colorado potato beetle, Leptinotarsa decemlineata, was studied by chronic feeding for 18 days on prey loaded with 1–16 μg OCI/day. Mortality of treated females was negligible, but fertility was reduced by up to 50%. Additional dose‐dependent effects in reproducing females included delayed oviposition, reduced fecundity, lower egg mass size, and reduced egg eclosion incidence. Females fed for 18 days on OCI at ≤4 μg/day returned to normal oviposition when switched to prey without OCI after 18 days of treatment, but negative effects persisted for at least 10 days at higher doses. Affected reproduction in P. bioculatus is consistent with the use of OCI‐sensitive digestive proteinases by this stinkbug. However, azocaseinase activity in whole body extracts of OCI‐fed females increased about twofold indicating compensation, and OCI‐sensitive proteinases were still present in extracts. When timed for delay to trigger attack on Colorado potato beetle larvae under controlled conditions, stinkbugs feeding on OCI appeared consistently hungrier than controls fed at similar rate, suggesting that predation by stinkbugs exposed to OCI‐recombinant foliage would be higher than normal. Arch. Insect Biochem. Physiol. 38:74–83, 1998. © 1998 Wiley‐Liss, Inc.

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16For parasitoids, the host represents the sole source of nutrients for the developing immature. 17Subsequently, host quality is an important factor affecting immature development and the 18 resulting fitness of the emerging parasitoid, with impacts on fecundity, longevity and offspring 19 sex ratio. Host age is an integral component of host quality and a key factor in host selection by 20 the female parasitoid. The current study aimed to investigate the effect of decreasing host quality 21 (determined by increasing host age) on adult life history traits (size, wing loading, longevity, and

Life tables of twospotted spider mite, Tetranychus urticae Koch (Acari: Tetranychidae), were studied on leaf discs and whole leaves of the common bean plant, Phaseolus vulgaris L. Data were analyzed based on the age-stage, two-sex life table theory. Durations of deutonymph stage, total preadult stage, and preoviposition period of mites reared on whole leaves (1.22, 7.6, and 0.29 d, respectively) were significantly shorter than those reared on leaf discs (1.73, 8.2, and 0.89 d, respectively). The lifetime fecundities were 22.81 and 12.05 offspring on whole leaves and leaf discs, respectively. Although hooked trichomes on the lower surface of the bean leaf contribute to adult mortality, spider mites successfully survive and reproduce on bean plants. The intrinsic rate of increase on whole leaves (0.235 d(-1)) was higher than that on leaf discs (0.159 d(-1). Higher intrinsic rate and fecundity found in mites reared on whole leaves may be due to the higher nutritional quality of whole leaves over leaf discs. We suggest that whole leaves should be used in future studies to better simulate realistic life history characteristics. The advantages of using age-stage, two-sex life tables over female age-specific life tables are discussed.