Jacqueline B. F. Geervliet

Comparative analysis of headspace volatiles from different caterpillar-infested or uninfested food plants of Pieris species

Plants that are infested by herbivores emit volatile cues that can be used by the natural enemies of the herbivores in their search for hosts. Based on results from behavioral studies, we investigated to what extent intact and herbivore-infested plant species and varieties from the food plant range of Pieris herbivore species differ in the composition of the volatile blends. Parasitoids of Pieris species, Cotesia glomerata and C. rubecula, show differential responses towards various herbivore-infested food plants, whereas differences in responses to plants infested by other herbivore species were less clear. Chemical analysis of the headspace samples of red cabbage, white cabbage, and nasturtium plants that were infested by P. brassicae or P. rapae larvae, or that were intact, yielded 88 compounds including alcohols, ketones, aldehydes, esters, nitriles, terpenoids, sulfides, (iso)thiocyanates, carboxylic acids, and others. The analysis revealed that herbivore-infested plants emit the largest number of compounds in the highest amounts. The plant species affected the volatile blend more than did the herbivore species, and differences between plant varieties were less pronounced than differences between plant species. Differences in headspace composition between plants infested by P. brassicae or P. rapae were mainly of a quantitative nature. Herbivore-infested nasturtium differed considerably from the cabbage varieties in a qualitative way. Headspace compositions of red and white cabbage varieties were comparable to that of the food plant Brussels sprouts (Brassica oleracea gemmifera cv. Titurel) as determined in earlier studies in our laboratory. With respect to plant response to herbivory, nasturtium differed considerably from the cabbage varieties analyzed so far and shows resemblance with Lima bean, cucumber, and corn. These plant species produce a greater quantity and variety of volatiles under herbivore attack than intact plants. The results of this study are discussed in relation to behavioral observations on C. glomerata and C. rubecula.

Volatiles from damaged plants as major cues in long‐range host‐searching by the specialist parasitoid Cotesia rubecula

The role of volatile stimuli in the long‐range host‐searching behaviour of the specialist parasitoid Cotesia rubecula Marshall (Hymenoptera: Braconidae) was studied. Components from the plant‐host‐complex Brussels sprouts (Brassica oleracea L. var. gemmifera (DC.) Schulz. cv. ‘Titurel’)‐Pieris rapae L. (Lepidoptera: Pieridae) were compared for their attractiveness in dual choice tests in a windtunnel. Stimuli from cabbage plants that were mechanically damaged or damaged by P. rapae caterpillars were more attractive to this parasitoid species than stimuli emitted by the host larvae or their faeces. Parasitoids preferred leaves from the plant‐host‐complex over artificially damaged leaves. Undamaged cabbage plants were the least attractive to the foraging females. These results indicate that in‐flight searching behaviour of C. rubecula is guided by plant‐derived information and that for this specialist species more reliable and specific host‐derived cues play a minor role at longer distances.

Learning to discriminate between infochemicals from different plant-host complexes by the parasitoids Cotesia glomerata and C. rubecula

Comparison of closely related species can elucidate adaptive differences in species characteristics. The present study compares the effect of experience on the host‐finding behaviour of two Cotesia (Hymenoptera: Braconidae) parasitoid species that differ in their degree of specialization. After multiple experiences with host larvae, females of the generalist parasitoid Cotesia glomerata showed a clear preference for volatiles from Pieris brassicae‐infested Brussels sprouts leaves over P. rapae‐infested Brussels sprouts leaves in two‐choice tests (‘preference learning at herbivore level’). A single experience with a host did not lead to such preferences.

Innate responses of the parasitoids Cotesia glomerata and C. rubecula (Hymenoptera: Braconidae) to volatiles from different plant-herbivore complexes.

To determine and compare innate preferences of the parasitoid speciesCotesia glomerata andC. rubecula for different plant-herbivore complexes, long-range (1-m) foraging behavior was studied in dual-choice experiments in a wind tunnel. In this study we tested the hypothesis that naive females of the specialistC. rubecula should show more pronounced preferences for different plant-herbivore complexes than females of the generalistC. glomerata. The herbivore species used were the pieridsPieris brassicae, P. rapae, P. Napi, andAporia crataegi and the nonhostsPlutella xylostella andMamestra brassicae. All herbivore species feed mainly on cabbage and wild crucifers, exceptAporia crataegi, which feeds on species of Rosaceae. Both parasitoid species preferred herbivore-damaged plants over nondamaged plants. NeitherC. rubecula norC. glomerata discriminated between plants infested by different caterpillar species, not even between plants infested by host-and nonhost species. Both parasitoid species showed preferences for certain cabbage cultivars and plant species. No differences were found in innate host-searching behavior betweenC. glomerata andC. rubecula. The tritrophic system cabbage-caterpillars-Cotesia sp. seems to lack specificity on the herbivore level, whereas on the plant level differences in attractiveness to parasitoids were found.

The role of host species, age and defensive behaviour on ovipositional decisions in a solitary specialist and gregarious generalist parasitoid (Cotesia species).

The main objective of this study was to determine the extent to which host acceptance behaviour as related to host species, age, and defensive behaviour might explain the differences in host use that exist between two congeneric and sympatric species of parasitic wasps. Cotesia glomerata (L.) (Hymenoptera: Braconidae) is gregarious and generalist on several species of Pieridae, whereas C. rubecula (Marshall) is solitary and specific to Pieris rapae (L.). Cotesia species differed in their responses to host species (P. brassicae (L.), P. napi (L.) and P. rapae) and developmental stage (early and late 1st, 2nd and 3rd instars). In no‐choice tests, host acceptance by C. rubecula was higher for P. rapae and females did not distinguish among the 6 host ages. In contrast, when foraging for P. brassicae and P. napi, C. rubecula females more readily attacked early first instar. Cotesia glomerata showed a higher degree of behavioural plasticity towards acceptance of Pieris host species and host age than did C. rubecula. Cotesia glomerata females parasitized the three Pieris species and showed higher acceptance of first and second instars over third instar. Oviposition success was also influenced by host defensive behaviour. The frequency and the effectiveness of defensive behaviour rose with increasing age of the host, P. brassicae being the most aggressive Pieris species. Furthermore, the mean duration of C. glomerata oviposition was significantly reduced by the defensive reactions of P. brassicae, which would likely affect parasitoid fitness as oviposition time is positively correlated to clutch size in C. glomerata. Acceptance frequencies corresponded well to field reports of Pieris‐Cotesia associations and to patterns of parasitoid larval performance, suggesting that the acceptance phase might be used as a reliable indicator of Cotesia host‐specificity.

Coexistence and niche segregation by field populations of the parasitoids Cotesia glomerata and C. rubecula in the Netherlands: predicting field performance from laboratory data.

Abstract Field experiments with foraging parasitoids are essential to validate the conclusions from laboratory studies and to interpret differences in searching and host selection behaviour of parasitoid species. Furthermore, field experiments can indicate whether the parameters measured in the laboratory are relevant to elucidation of the ecological processes under study, such as adaptation or species interactions. In previous extensive laboratory studies we studied plant- and host-searching behaviour, host acceptance, host suitability; host plant preference, and learning of two congeneric parasitoids of Pieris caterpillars: the generalist Cotesia glomerata, which has been reported to attack several Pieridae species, and C. rubecula, a specialist of the small cabbage white Pieris rapae. In the present field study our aim was to verify the importance of these previous laboratory findings for explaining the performance of these two species in the field. We investigated experimentally whether parasitism on three Pieris species varied with parasitoid species and with food plant of the caterpillars. We exposed different types of host plants, infested with different Pieris species, to parasitism by natural populations of Cotesia species, by setting the experimental plants out in Brussels sprouts cabbage fields. Furthermore we made direct observations of parasitoid foraging in the field. In general, the field results confirmed our predictions on the range of host plant and host species used in the field. The two Cotesia species appear to coexist through niche segregation, since C. glomerata was mainly recovered from P. brassicae and C. rubecula from P. rapae. Although C. glomerata is a generalist at the species level, it can be a specialist at the population level under certain ecological circumstances. Our study shows the importance of variation in host plant attraction and host species acceptance in restricting host plant and host diet in the field. Furthermore the results suggest that, at least in the Netherlands, specialisation of C. glomerata on P. brassicae may occur as a result of C. rubecula outcompeting C. glomerata in P. rapae larvae.

Effects of Pieris host species on life history parameters in a solitary specialist and gregarious generalist parasitoid (Cotesia species)

Host specificity and host selection by insect parasitoids are hypothesized to be correlated with suitability of the hosts for parasitoid development. The present study investigates the correlation between host suitability and earlier studied host‐finding behaviour of two closely related braconid larval parasitoid species, the generalist Cotesia glomerata (L.) and the specialist C. rubecula (Marshall) (Hymenoptera: Braconidae). We compared the capability of both parasitoid species to parasitize and develop in three Pieris host species, i.e. P. brassicae (L.), P. rapae (L.) and P. napi (L.) (Lepidoptera: Pieridae). In laboratory experiments, we measured the effect of host species on fitness parameters such as survival, development, sex ratio and size of parasitoid progeny. The results show that C. glomerata is capable of developing in the three host species, with significant differences in parasitoid survival, clutch size and adult weight among Pieris species. The host range for development was more restricted for C. rubecula. Although C. rubecula is physiologically able to develop in P. brassicae larvae, parasitoid fitness is negatively affected by this host species, compared to its most regular host, P. rapae. A comparison of the present data on host suitability with earlier studies on host‐searching behaviour suggests that the host‐foraging behaviour of both parasitoid species not only leads to selection of the most suitable host species for parasitoid development, but also plays a significant role in shaping parasitoid host range.