Parasitoid Cotesia Rubecula Research Articles

Innate and learned olfactory attraction to flowering plants by the parasitoid Cotesia rubecula (Marshall, 1885) (Hymenoptera: Braconidae): Potential impacts on conservation biological control

In conservation biological control, flowers can be used to increase the biological control potential of parasitoids, which benefit from the offered food sources. Besides exhibiting exploitable nectar, flowers should preferably be olfactorily attractive, as highly attractive flowers are easily located, reducing the time spent searching for food and subsequently increasing the per capita host searching efficiency. In this study we thus focused on the olfactory attractiveness of Fagopyrum esculentum Moench (Polygonaceae), Centaurea cyanus L. (Asteraceae) and Vicia sativa L. (Fabaceae) to Cotesia rubecula (Marshall, 1885) (Hymenoptera: Braconidae), a larval parasitoid of the cabbage pest Pieris rapae (Linnaeus, 1758) (Lepidoptera: Pieridae). With a Y-tube olfactometer we found that C. cyanus and to a lesser extent V. sativa successfully attract C. rubecula. Also F. esculentum attracts C. rubecula, but only after a rewarding feeding experience. All three tested flowers seem to be suitable to be exploited in conservation biological control programs to control P. rapae in brassica fields. Even though not every flower offering accessible nectar is also innately attractive, it can still be suitable for conservation biological control purposes as feeding experience can change this attraction. In fact, the application of mixtures containing attractive and rewarding flowers could help increase the success of such programs.

Effect ofCotesia rubecula(Hymenoptera: Braconidae) on Survival of Larval Cohorts ofPieris rapae(Lepidoptera: Pieridae) on Collards: Evaluation of an Introduced Biological Control Agent

Survival of artificial cohorts of larvae of the imported cabbageworm, Pieris rapae (L.), was assessed in a collard (Brassica oleracea L., var. acephala) patch on an organic vegetable farm in western Massachusetts. We recorded a significant drop in survival of larvae between the 4th and 5th instar due to parasitism by the parasitoid Cotesia rubecula (Marshall), which was introduced to New England from China in 1988. This was a large change from the survivorship pattern of P. rapae observed in the same area in 1985–1986, in which there was a significant drop in survival between 5th instars and pupae due to Cotesia glomerata (L.), the dominant parasitoid of P. rapae in New England at the time. In our study, parasitism by C. rubecula averaged 62.4 ± 8.6%, while no parasitism by C. glomerata was observed. The introduction of C. rubecula is now very likely to be benefitting organic vegetable producers in western Massachusetts, killing more larvae and at an earlier life stage than the previous parasitoid, reducing the potential for damage to cole crops.

Smelling the Wood from the Trees: Non-Linear Parasitoid Responses to Volatile Attractants Produced by Wild and Cultivated Cabbage

Despite a large number of studies on herbivore-induced plant volatiles (HIPVs), little is known about which specific compounds are used by natural enemies to locate prey- or host- infested plants. In addition, the role of HIPVs in attracting natural enemies has been restricted largely to agricultural systems. Differences in volatile blends emitted by cultivars and plants that originate from wild populations may be attributed to potentially contrasting selection regimes: natural selection among the wild types and artificial selection among cultivars. A more realistic understanding of these interactions in a broader ecological and evolutionary framework should include studies that involve insect herbivores, parasitoids, and wild plants on which they naturally interact in the field. We compared the attractiveness of HIPVs emitted by wild and cultivated cabbage to the parasitoid Cotesia rubecula, and determined the chemical composition of the HIPV blends to elucidate which compounds are involved in parasitoid attraction. Wild and cultivated cabbage differed significantly in their volatile emissions. Cotesia rubecula was differentially attracted to the wild cabbage populations and preferred wild over cultivated cabbage. Isothiocyanates, which were only emitted by the wild cabbages, may be the key components that explain the preference for wild over cultivated cabbage, whereas terpenes may be important for the differential attraction among the wild populations. Volatile analysis revealed that parasitoid attraction cannot be explained by simple linear relationships. Our results suggest that unraveling which compound(s) are innately attractive to parasitoids of cabbage pests should include wild Brassicaceae.

Influences of plant species on life history traits of Cotesia rubecula (Hymenoptera: Braconidae) and its host Pieris rapae (Lepidotera: Pieridae)

The relative suitability of four plants was studied for larvae of Pieris rapae L. and its parasitoid Cotesia rubecula (Marshall). For unparasitized P. rapae, pupal dry weight and egg-pupa growth rate were higher on cabbage, radish and nasturtium than on Indian hedge mustard. Larval developmental rate and size were greatest for C. rubecula when its host was feeding on nasturtium. Wasp survival was not affected by the host insect/plant combination in which the parasitoid developed. These results indicate that the plant on which host larvae feed is an important factor in development of the parasitoid.

Formation of Simple Nitriles upon Glucosinolate Hydrolysis Affects Direct and Indirect Defense Against the Specialist Herbivore, Pieris rapae

The glucosinolate-myrosinase system, found in plants of the order Brassicales, has long been considered an effective defense system against herbivores. The defensive potential of glucosinolates is mainly due to the products formed after myrosinase-catalyzed hydrolysis upon tissue damage. The most prominent hydrolysis products, the isothiocyanates, are toxic to a wide range of organisms, including herbivorous lepidopterans. In contrast, little is known about the biological activities of alternative hydrolysis products such as simple nitriles and epithionitriles that are formed at the expense of isothiocyanates in the presence of epithiospecifier proteins (ESPs). Here, we used transgenic Arabidopsis thaliana (Brassicaceae) plants overexpressing ESP (35S:ESP plants) to investigate the effects of simple nitriles on direct and indirect defense against the specialist cabbage white butterfly Pieris rapae L. (Lepidoptera, Pieridae). In the 35S:ESP plants, glucosinolates are hydrolyzed mainly to simple nitriles upon tissue disruption, while isothiocyanates are the predominant hydrolysis products in Columbia-0 (Col-0) wild-type plants. The parasitoid Cotesia rubecula (Hymenoptera, Braconidae), a specialist on P. rapae larvae, was significantly more attracted to P. rapae-infested 35S:ESP plants than to P. rapae-infested Col-0 wild-type plants in a wind tunnel setup. Furthermore, female P. rapae butterflies laid more eggs on Col-0 wild-type plants than on 35S:ESP plants when the plants had been damaged previously. However, when given a choice to feed on 35S:ESP or Col-0 plants, caterpillars did not discriminate between the two genotypes. Growth rate and developmental time were not significantly different between caterpillars that were reared on 35S:ESP or Col-0 plants. Thus, the production of simple nitriles instead of isothiocyanates, as catalyzed by ESP, can promote both direct and indirect defense against the specialist herbivore P. rapae.

Herbivore-Induced Plant Volatiles Mediate In-Flight Host Discrimination by Parasitoids

Herbivore feeding induces plants to emit volatiles that are detectable and reliable cues for foraging parasitoids, which allows them to perform oriented host searching. We investigated whether these plant volatiles play a role in avoiding parasitoid competition by discriminating parasitized from unparasitized hosts in flight. In a wind tunnel set-up, we used mechanically damaged plants treated with regurgitant containing elicitors to simulate and standardize herbivore feeding. The solitary parasitoid Cotesia rubecula discriminated among volatile blends from Brussels sprouts plants treated with regurgitant of unparasitized Pieris rapae or P. brassicae caterpillars over blends emitted by plants treated with regurgitant of parasitized caterpillars. The gregarious Cotesia glomerata discriminated between volatiles induced by regurgitant from parasitized and unparasitized caterpillars of its major host species, P. brassicae. Gas chromatography-mass spectrometry analysis of headspace odors revealed that cabbage plants treated with regurgitant of parasitized P. brassicae caterpillars emitted lower amounts of volatiles than plants treated with unparasitized caterpillars. We demonstrate (1) that parasitoids can detect, in flight, whether their hosts contain competitors, and (2) that plants reduce the production of specific herbivore-induced volatiles after a successful recruitment of their bodyguards. As the induced volatiles bear biosynthetic and ecological costs to plants, downregulation of their production has adaptive value. These findings add a new level of intricacy to plant-parasitoid interactions.

Isolation of an imaginal disc growth factor homologue from Pierisrapae and its expression following parasitization by Cotesiarubecula

Endoparasitoid insects introduce maternal factors into the body of their host at oviposition to suppress cellular defences for the protection of the developing parasitoid. We have shown that transient expression of polydnavirus genes from a hymenopteran parasitoid Cotesia rubecula (CrPDV) is responsible for the inactivation of hemocytes from the lepidopteran host Pieris rapae. Since the observed downregulation of CrPDV genes in infected host tissues is not due to cis-regulatory elements at the CrV1 gene locus, we speculated that the termination of CrPDV gene expression may be due to cellular inactivation caused by the CrV1-mediated immune suppression of infected tissues. To test this assumption, we isolated an imaginal disc growth factor (IDGF) that is expressed in fat body and hemocytes, the target of viral infection and expression of CrPDV genes. Time-course experiments showed that the level of P. rapae IDGF is not affected by parasitization and polydnavirus infection. However, the amount of highly expressed genes, such as storage proteins, arylphorin and lipophorin, are significantly reduced following parasitization.

Signal transduction downstream of salicylic and jasmonic acid in herbivory‐induced parasitoid attraction by Arabidopsis is independent of JAR1 and NPR1

ABSTRACTPlants can defend themselves indirectly against herbivores by emitting a volatile blend upon herbivory that attracts the natural enemies of these herbivores, either predators or parasitoids. Although signal transduction in plants from herbivory to induced volatile production depends on jasmonic acid (JA) and salicylic acid (SA), the pathways downstream of JA and SA are unknown. Use of Arabidopsis provides a unique possibility to study signal transduction by use of signalling mutants, which so far has not been exploited in studies on indirect plant defence. In the present study it was demonstrated that jar1‐1 and npr1‐1 mutants are not affected in caterpillar (Pieris rapae)‐induced attraction of the parasitoid Cotesia rubecula. Both JAR1 and NPR1 (also known as NIM1) are involved in signalling downstream of JA in induced defence against pathogens such as induced systemic resistance (ISR). NPR1 is also involved in signalling downstream of SA in defence against pathogens such as systemic acquired resistance (SAR). These results demonstrate that signalling downstream of JA and SA differs between induced indirect defence against herbivores and defence against pathogens such as SAR and ISR. Furthermore, it was demonstrated that herbivore‐derived elicitors are involved in induced attraction of the parasitoid Cotesia rubecula

Attraction of the specialist parasitoid Cotesia rubecula to Arabidopsis thaliana infested by host or non‐host herbivore species

AbstractIn this study we investigated whether in a two‐choice set‐up the parasitoid Cotesia rubecula (Marshall) (Hymenoptera, Braconidae) distinguishes between volatiles emitted by Arabidopsis thaliana (L.) Heynh. (Brassicaceae) infested with its host, Pieris rapae (L.) (Lepidoptera: Pieridae) and Arabidopsis infested with non‐host herbivores. Four non‐host herbivore species were tested: the caterpillars Plutella xylostella (L.) (Lepidoptera: Plutellidae) and Spodoptera exigua (Hübner) (Lepidoptera: Noctuidae), both chewing insects, the spider mite Tetranychus urticae (Koch) (Acari: Tetranychidae), which punctures parenchymal cells, and the aphid Myzus persicae (Sulzer) (Hemiptera: Aphidoidea), which is a phloem‐feeder. Compared with undamaged plants, C. rubecula females were more attracted to Arabidopsis plants infested by P. rapae, P. xylostella, S. exigua, or T. urticae, but not to plants infested by M. persicae. The parasitoids preferred host‐infested plants to spider mite‐ or aphid‐infested plants, but not to plants infested with non‐host caterpillars (P. xylostella or S. exigua). The data show that when Arabidopsis plants are infested with a leaf tissue‐damaging herbivore they emit a volatile blend that attracts C. rubecula females and the wasps only discriminate between a host and non‐host herbivore when the type of damage is different (chewing vs. piercing). When Arabidopsis is infested with a herbivore that hardly damages leaf tissue, C. rubecula females are not attracted. These results may be explained by differences in the amount of damage and in the relative importance of different signal‐transduction pathways induced by different types of herbivores.

Introduced braconid parasitoids and range reduction of a native butterfly in New England

Abstract Pieris napi oleracea Harris is a native pierid butterfly that has suffered a range reduction in New England that began after the invasion of its range by the non-native congener Pieris rapae L. and one of its braconid parasitoids, Cotesia glomerata (L.). P. napi has nearly disappeared from Massachusetts, but remains common in northern Vermont. We investigated food plant abundance and Cotesia spp. larval parasitism as possible factors to explain the historical changes in P. napi ’s distribution. We found that the current range of P. napi was not explained by the abundance of its key first generation food plant (two-leafed toothwort, Cardamine diphylla [Michx.]). We also found that levels of Cotesia spp. parasitism in meadows in the second generation were similar in Vermont and Massachusetts. Further, we found that both C. glomerata and the related introduced Pieris spp. parasitoid Cotesia rubecula (Marshall) forage for hosts predominantly in sunny meadows and not in woods, where the first generation of P. napi occurs. We found that under field conditions in meadow habitats, C. glomerata parasitizes P. napi at higher rates than P. rapae . We postulate that the persistence of P. napi in Vermont and its disappearance in Massachusetts is caused by high parasitism of the second generation by C. glomerata in meadow habitats, coupled with a north–south cline in the rate of commitment of first generation P. napi pupae to diapause, such that northern populations act functionally as univoltine species developing in a parasitoid free habitat (woods), while southern populations acted as a bivoltine species and went extinct due to low survival in the second generation in meadows due to C. glomerata parasitism.

Induced parasitoid attraction by Arabidopsis thaliana: involvement of the octadecanoid and the salicylic acid pathway.

Plants can use indirect defence mechanisms to protect themselves against herbivorous insects. An example of such an indirect defence mechanism is the emission of volatiles by plants induced by herbivore feeding. These volatiles can attract the natural enemies of these herbivores, for example, parasitoid wasps. Here, it is shown that the octadecanoid and the salicylic acid pathways are involved in the induced attraction of the parasitoid wasp Cotesia rubecula by Arabidopsis thaliana infested with the herbivore Pieris rapae. Besides exogenous application of jasmonic acid or salicylic acid, use is also made of transgenic Arabidopsis that do not show induced jasmonic acid levels after wounding (S-12) and transgenic Arabidopsis that do not accumulate salicylic acid (NahG). Treatment of Arabidopsis with jasmonic acid resulted in an increased attraction of parasitoid wasps compared with untreated plants, whereas treatment with salicylic acid did not. Transgenic plants impaired in the octadecanoid or the salicylic acid pathway were less attractive than wild-type plants.

Feeding and survival in parasitic wasps: sugar concentration and timing matter

Sugar consumption can increase the longevity and lifetime fecundity of many species of parasitic wasps. Consequently, for these insects the availability of sugar sources in the field is important for their reproductive success. As sugar sources can be highly variable in quantity, space and time, the chances of finding a sufficient amount of sugar to increase longevity might be very low. Therefore, the reward from a single feeding event can be critical for the forager's fitness. We measured the longevity of the parasitoid Cotesia rubecula after a single honey meal differing in sugar concentration (25, 47, 86% w/w) and timing (day of emergence and 24 h later). Survival was analysed with Cox's Proportional Hazards Model. The risk of starving to death in sugar‐fed wasps was reduced by 0–73% in comparison to unfed wasps, depending on sugar concentration and timing. Longevity was significantly increased by sugar concentration and by feeding later in life. Our results suggest that in the field, adult C. rubecula has to locate food at least once per day to avoid starvation.

The Effect of Resource Aggregation at Different Scales: Optimal Foraging Behavior ofCotesia rubecula

Resources can be aggregated both within and between patches. In this article, we examine how aggregation at these different scales influences the behavior and performance of foragers. We developed an optimal foraging model of the foraging behavior of the parasitoid wasp Cotesia rubecula parasitizing the larvae of the cabbage butterfly Pieris rapae. The optimal behavior was found using stochastic dynamic programming. The most interesting and novel result is that the effect of resource aggregation within and between patches depends on the degree of aggregation both within and between patches as well as on the local host density in the occupied patch, but lifetime reproductive success depends only on aggregation within patches. Our findings have profound implications for the way in which we measure heterogeneity at different scales and model the response of organisms to spatial heterogeneity.

The Effect of Resource Aggregation at Different Scales: Optimal Foraging Behavior of Cotesia rubecula

Resources can be aggregated both within and between patches. In this article, we examine how aggregation at these different scales influences the behavior and performance of foragers. We developed an optimal foraging model of the foraging behavior of the parasitoid wasp Cotesia rubecula parasitizing the larvae of the cabbage butterfly Pieris rapae. The optimal behavior was found using stochastic dynamic programming. The most interesting and novel result is that the effect of resource aggregation within and between patches depends on the degree of aggregation both within and between patches as well as on the local host density in the occupied patch, but lifetime reproductive success depends only on aggregation within patches. Our findings have profound implications for the way in which we measure heterogeneity at different scales and model the response of organisms to spatial heterogeneity.

Herbivore-induced volatile production by Arabidopsis thaliana leads to attraction of the parasitoid Cotesia rubecula: chemical, behavioral, and gene-expression analysis.

Many plant species defend themselves against herbivorous insects indirectly by producing volatiles in response to herbivory. These volatiles attract carnivorous enemies of the herbivores. Research on the model plant Arabidopsis thaliana (L.) Heynh. has contributed considerably to the unraveling of signal transduction pathways involved in direct plant defense mechanisms against pathogens. Here, we demonstrate that Arabidopsis is also a good candidate for studying signal transduction pathways involved in indirect defense mechanisms by showing that: (1) Adult females of Cotesia rubecula, a specialist parasitic wasp of Pieris rapae caterpillars, are attracted to P. rapae-infested Arabidopsis plants. (2) Arabidopsis infested by P. rapae emits volatiles from several major biosynthetic pathways, including terpenoids and green leaf volatiles. The blends from herbivore-infested and artificially damaged plants are similar. However, differences can be found with respect to a few components of the blend, such as two nitriles and the monoterpene myrcene, that were produced exclusively by caterpillar-infested plants, and methyl salicylate, that was produced in larger amounts by caterpillar-infested plants. (3) Genes from major biosynthetic pathways involved in volatile production are induced by caterpillar feeding. These include AtTPS10, encoding a terpene synthase involved in myrcene production, AtPAL1, encoding phenylalanine ammonia-lyase involved in methyl salicylate production, and AtLOX2 and AtHPL, encoding lipoxygenase and hydroperoxide lyase, respectively, both involved in the production of green leaf volatiles. AtAOS, encoding allene oxide synthase, involved in the production of jasmonic acid, also was induced by herbivory.

Role of volatile inforchemicals emitted by feces of larvae in host-searching behavior of parasitoidCotesia rubecula (Hymenoptera: Braconidae): A behavioral and chemical study

The role of volatile infochemicals emitted by feces of larvae in the host-searching behavior of the parasitoidCotesia rubecula was evaluated during single- and dual-choice tests inside a wind tunnel. The following treatments were tested: feces produced by second and fourth instars ofPieris rapae (preferred host), second instars ofP. brassicae (inferior host), second instars ofP. napi (nonhost), and wet feces of second instars ofP. rapae. During a single-choice situation females ofC. rubecula oriented to all types of feces tested. When a preference was to be made,C. rubecula preferred feces of second instars ofP. rapae over that of fourth, feces ofP. rapae over that ofP. brassicae, feces ofP. napi over that ofP. brassicae, and wet over normal host feces. No preference was exhibited between feces of second instars ofP. napi and that of second instars ofP. rapae. The relative importance of infochemicals from host feces versus plant damage caused by host larvae to the searching behavior ofC. rubecula was also evaluated. Plant damage was more important to the searching females than host feces when feces were present in specific concentrations in relation to damage. The volatiles released by normal and wet feces of second instars ofP. rapae, wet feces of fourth instars ofP. rapae, and normal and wet feces ofP. brassicae were collected and identified. Overall, 85 chemical compounds were recorded belonging to the following chemical groups: alcohols, ketones, aldehydes, esters, isothiocyanates, sulfides, nitriles, furanoids, terpenoids and pyridines. The blend of chemicals emitted by feces of different instars ofP. rapae and different species ofPieris exhibited an instar and species specificity in both quantity and quality. Wetting of normal feces increased the amount of volatile chemicals released, and it was also responsible for the appearance of new compounds. The role of feces of larvae in the host-seeking behavior ofC. rubecula is discussed.

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

AbstractThe 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.

The effect of food deprivation on the innate visual and olfactory preferences in the parasitoid Cotesia rubecula

The response of Cotesia rubecula (Hymenoptera: Braconidae) to olfactory and visual flower stimuli was tested. It was demonstrated that parasitoids can use both flower-odors and -colors during food foraging. The response of parasitoids to food-indicating stimuli depended on the hunger state of the individual. Given a choice in a Y-tube olfactometer between flower odors and odors from host-infested leaves, food-deprived individuals chose flower odors, while sugar-fed individuals preferred host associated odors. In flight chamber experiments, food-deprived parasitoids landed more often and spent more time searching on yellow targets, while sugar-fed individuals displayed a higher overall foraging activity, without reacting differentially to yellow.

Plant-natural enemy association in tritrophic system,Cotesia rubecula-Pieris rapae-brassicaceae (Cruciferae). III: Collection and identification of plant and frass volatiles

To elucidate the identity of the volatile compounds that could be involved in the searching behavior of the parasitoidCotesia rubecula Marshall (Hymenoptera: Braconidae), the volatiles released by cabbage and frass of Lepidoptera feeding on cabbage were collected and analyzed using a gas chromatograph-mass spectrometer. The volatiles emitted by intact cabbage were α-pinene, β-pinene, myrcene, 1,8-cineole,n-hexyl acetate,cis-3-hexen-1-yl acetate, and dimethyl trisulfide. Mechanical damage on an intact plant induced the release of two more compounds,trans-2-hexenal and 1-methoxy-3-methylene-2-pentanone. Current feeding by larvae ofPieris rapae L. (Pieridae) induced the plant to release all the compounds released after mechanical damage and additionally 4-methyl-3-pentenal and allyl isothiocyanate. Current feeding by larvae ofPlutella xylostella L. (Plutellidae) induced the plant to release all the compounds present after mechanical damage and additionally allyl isothiocyanate. The volatiles emitted after feeding by the lepidopterans had ceased were the same as those emitted by cabbage damaged by mechanical means. The blend of volatiles emitted by frass was comprised of plant chemicals, mainly sulfur compounds. Frass ofP. rapae emitted allyl isothiocyanate, methyl isothiocyanate, methyl propyl sulfide, dimethyl trisulfide,S-methyl methane thiosulfinate, 4-methyl-3-pentenal,trans-2-hexenal, and 2,3-dihydro-4-methyl furan. Frass ofP. xylostella emitted only dimethyl trisulfide andS-methyl methane thiosulfinate. The blend of volatiles emitted by frass is herbivore-species specific.

Plant-natural enemy association in the tritrophic system,Cotesia rubecula-Pieris rapae-brassiceae (cruciferae): I. Sources of infochemicals

The role of airborne infochemicals in host selection by the parasitoidCotesia rubecula (Marshal) (Hymenoptera: Braconidae) was examined in a wind tunnel. To elucidate the role of volatile chemicals in attractingC. rubecula to cabbage infested by the host [Pieris rapae L. (Lepidoptera: Pieridae)], the potential sources of volatiles related toP. rapae infestation on cabbage were tested individually. The responses of females to nonhost plant species, bean and geranium, as well as to frass of a nonhost lepidopteran were also examined.C. rubecula was attracted to cabbage previously infested byP. rapae and to frass and regurgitate ofP. rapae. No attraction was observed to larvae ofP. rapae alone. Females were also attracted to mechanically damaged cabbage, cabbage previously infested byPlutella xylostella L. (Lepidoptera: Plutellidae) (a nonhost lepidopteran herbivore), and cabbage previously infested by snails (a nonhost, noninsect herbivore). Intact cabbage, bean, and geranium plants elicited no attraction. A low frequency of attraction was observed to mechanically damaged bean and geranium. Attraction was also observed to frass ofP. xylostella. Volatiles from cabbage related to damage, and volatiles from frass and regurgitate of the host seem to play an important role in guidingC. rubecula to plants infested by its host.