Plant-natural enemy association in the tritrophic systemCotesia rubecula-Pieris rapae-brassicaceae (cruciferae): II. Preference ofC. rubecula for landing and searching

The responses of the parasitoidCotesia rubecula to differently damaged cabbages were recorded during a series of choice tests. To determine if flyingC. rubecula can discriminate differences in the blend of volatiles emitted by cabbages damaged by different causes and how plant volatiles released from a distant source affect the searching behavior ofC. rubecula once searching on a plant, wasps were presented with a choice of plants located one behind the other and separated by a distance of 15 cm. The sources of damage were: cabbage damaged by the host (Pieris rapae), by a nonhost lepidopteran herbivore (Plutella xylostella), by a nonhost, noninsect herbivore (snail), and by mechanical means. The results showed that the site of first landing and the time spent searching on the leaves was influenced by the type of damage inflicted on plants. Wasps preferred to land on cabbages damaged by host and nonhost species of Lepidoptera over those damaged by snails and mechanical means. No preference was observed for first landing between cabbages damaged by the two species of Lepidoptera or between cabbages damaged by snails and mechanical means. Cabbage damaged byP. rapae was searched most intensively, followed by cabbage damaged byP. xylostella, cabbage damaged by snails, and cabbage damaged by mechanical means.C. rubecula differentiates between the volatile blends emitted by differently damaged cabbages, and it is attracted to volatiles related to recent lepidopteran damage. Wasps searched longer on freshly damaged than on leaves with older damage.     

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 -pinene, -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.     

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.     

Effect of the Presence of a Nonhost Herbivore on the Response of the Aphid Parasitoid <Emphasis Type="Italic">Diaeretiella rapae</Emphasis> to Host-infested Cabbage Plants

The vast majority of studies of plant indirect defense strategies have considered simple tritrophic systems that involve plant responses to attack by a single herbivore species. However, responses by predators and parasitoids to specific, herbivore-induced, volatile blends could be compromised when two or more different herbivores are feeding on the same plant. In Y-tube olfactometer studies, we investigated the responses of an aphid parasitoid, Diaeretiella rapae (McIntosh) (Hymenoptera: Braconidae), to odors from cabbage plants infested with the peach-potato aphid Myzus persicae (Sulzer) (Homoptera: Aphididae), in both the presence and absence of a lepidopteran caterpillar, Plutella xylostella L. (Lepidoptera: Plutellidae). Female parasitoids chose aphid-infested plants over uninfested plants but did not distinguish between caterpillar-infested and uninfested plants. When given a choice between odors from an aphid-infested plant and those from a plant infested with diamondback moth larvae, they significantly chose the former. Furthermore, the parasitoids responded equally to odors from a plant infested with aphids only and those from a plant infested with both aphids and caterpillars. The results support the hypothesis that the aphid and the caterpillar induce different changes in the volatile profile of cabbage plants and that D. rapae females readily distinguish between the two. Furthermore, the changes to the plant volatile profile induced by the caterpillar damage did not hinder the responses of the parasitoid to aphid-induced signals.     

Relative importance of semiochemicals from first and second trophic levels in host foraging behavior ofAphidius ervi

The responses of femaleAphidius ervi to odors from a host food plant (Vicia faba), host aphids (Acyrthosiphon pisum), nonhost aphids (Aphis fabae), and aphid-plant complexes were investigated in a specially designed wind tunnel and a Y-tube olfactometer. In single-target (no-choice) and two-target (dual-choice) experiments, plant volatiles played a crucial role in the host foraging behavior ofA. ervi. The odor from theA. pisum-plant complex elicited the strongest responses byA. ervi females, followed by the odor from plants previously damaged by the feeding ofA. pisum. There was a significantly weaker response to odor fromA. pisum in the absence of the plant and to undamaged plants. Similarly, mechanically damaged plants and plants infested with the nonhost aphidA. fabae did not elicit strong responses. A plant that had been damaged byA. pisum and subsequently washed with distilled water was as attractive as an unwashed, previously infested plant.Aphidius ervi probably overcomes the reliability-detectability problem by selectively responding to herbivore-induced, volatile, semiochemical cues emitted by the first trophic level and by distinguishing between the volatiles induced by host and nonhost aphids.     

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.     

Systemic release of herbivore-induced plant volatiles by turnips infested by concealed root-feeding larvae Delia radicum L

When attacked by herbivorous insects, many plants emit volatile compounds that are used as cues by predators and parasitoids foraging for prey or hosts. While such interactions have been demonstrated in several host–plant complexes, in most studies, the herbivores involved are leaf-feeding arthropods. We studied the long-range plant volatiles involved in host location in a system based on a very different interaction since the herbivore is a fly whose larvae feed on the roots of cole plants in the cabbage root fly, Delia radicum L. (Diptera: Anthomyiidae). The parasitoid studied is Trybliographa rapae Westwood (Hymenoptera: Figitidae), a specialist larval endoparasitoid of D. radicum. Using a four-arm olfactometer, the attraction of naive T. rapae females toward uninfested and infested turnip plants was investigated. T. rapae females were not attracted to volatiles emanating from uninfested plants, whether presented as whole plants, roots, or leaves. In contrast, they were highly attracted to volatiles emitted by roots infested with D. radicum larvae, by undamaged parts of infested roots, and by undamaged leaves of infested plants. The production of parasitoid-attracting volatiles appeared to be systemic in this particular tritrophic system. The possible factors triggering this volatile emission were also investigated. Volatiles from leaves of water-stressed plants and artificially damaged plants were not attractive to T. rapae females, while volatiles emitted by leaves of artificially damaged plants treated with crushed D. radicum larvae were highly attractive. However, T. rapae females were not attracted to volatiles emitted by artificially damaged plants treated only with crushed salivary glands from D. radicum larvae. These results demonstrate the systemic production of herbivore-induced volatiles in this host-plant complex. Although the emission of parasitoid attracting volatiles is induced by factors present in the herbivorous host, their exact origin remains unclear. The probable nature of the volatiles involved and the possible origin of the elicitor of volatiles release are discussed.     

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.     

Host-specific recognition kairomone for the parasitoidMicroplitis croceipes (Cresson)

A water-extractable host recognition kairomone in frass of corn earworm,Helicoverpa zea Boddie (Lepidoptera: Noctuidae), host larvae stimulates antennation by females of the parasitoidMicroplitis croceipes Cresson (Braconidae: Hymenoptera). In addition, when the wasps contact water extracts of host frass they will subsequently fly in a flight tunnel to odor associated with the extract. Contact with water extracts of cowpea leaves or with water extracts of frass from larvae of nonhost beet armyworm, fall armyworm, or cabbage looper that were fed cowpea leaves does not stimulate antennation, nor do wasps fly to associated odors after contact with these substances. However, contact with the water extract of host frass in association with hexane extract of cowpea-fed nonhost frass will induce the wasps to subsequently fly to the hexane extract of the nonhost frass when it is used as an odor source in a flight tunnel. Thus the host-specific kairomone by whichM. croceipes recognizes the frass of its host is extractable with water. This substance plays a crucial role in the foraging behavior of this parasitoid by allowing it to recognize host frass and to learn to search for odors originating from plants on which the host is feeding.     

Induction of parasitoid attracting synomone in brussels sprouts plants by feeding ofPieris brassicae larvae: Role of mechanical damage and herbivore elicitor

Induction of plant defense in response to herbivory includes the emission of synomones that attract the natural enemies of herbivores. We investigated whether mechanical damage to Brussels sprouts leaves (Brassica oleracea var.gemmifera) is sufficient to obtain attraction of the parasitoidCotesia glomerata or whether feeding byPieris brassicae caterpillars elicits the release of synomones not produced by mechanically damaged leaves. The response of the parasitoidCotesia glomerata to different types of simulated herbivory was observed. Flight-chamber dual-choice tests showed that mechanically damaged cabbage leaves were less attractive than herbivore-damaged leaves and mechanically damaged leaves treated with larval regurgitant. Chemical analysis of the headspace of undamaged, artificially damaged, caterpillar-infested, and caterpillar regurgitant-treated leaves showed that the plant responds to damage with an increased release of volatiles. Greenleaf volatiles and several terpenoids are the major components of cabbage leaf headspace. Terpenoids are emitted in analogous amounts in all treatments, including undamaged leaves. On the other hand, if the plant is infested by caterpillars or if caterpillar regurgitant is applied to damaged leaves, the emission of green-leaf volatiles is highly enhanced. Our data are in contrast with the induction of more specific synomones in other plant species, such as Lima bean and corn.     

Host microhabitat location by stem-borer parasitoidCotesia flavipes: the role of herbivore volatiles and locally and systemically induced plant volatiles

The origin of olfactory stimuli involved in the host microhabitat location inCotesia flavipes, a parasitoid of stem-borer larvae, was investigated in a Y-tube olfactometer. The response of femaleC. flavipes towards different components of the plant-host complex, consisting of a maize plant infested with two or more larvae of the stem borerChilo partellus, was tested in dualchoice tests. The concealed lifestyle of the stem-borer larvae did not limit the emission of volatiles attractive to a parasitoid. A major source of the attractive volatiles from the plant-host complex was the stem-borer-injured stem, including the frass produced by the feeding larvae. Moreover, the production of volatiles attractive to a parasitoid was not restricted to the infested stem part but occurs systemically throughout the plant. The uninfested leaves of a stem-borer-infested plant were found to emit volatiles that attract femaleC. flavipes. We further demonstrate that an exogenous elicitor of this systemic plant response is situated in the regurgitate of a stem-borer larva. When a minor amount of regurgitate is inoculated into the stem of an uninfested plant, the leaves of the treated plant emit volatiles that attract femaleC. flavipes.     

Secondary attraction inGnathotrichus retusus and cross-attraction ofG. sulcatus (Coleoptera: Scolytidae)

Laboratory bioassays indicated that frass produced byGnathotrichus retusus males and male gut extracts were attractive to both sexes of beetles. The frass became attractive to females within 2 days of the commencement of boring activity, and attractive frass was produced for at least lOdays. Attraction of frass rose sharply after pairing of males with females. However, this increase in attraction could be due to increased boring activity and pheromone production by males following establishment of the mutualistic fungus. In field trapping experiments, male-infested logs were always highly attractive, but in one experiment, female-infested logs and logs infested by both sexes were also attractive. Thus, females could be involved in secondary attraction. High cross-attraction ofG. sulcatus toG. retusus infested logs in field experiments, as well as a moderate but significant response byG. retusus to sulcatol in the laboratory, suggests that they share sulcatol as a common pheromone.Research supported by Canadian Forestry Service Science Subvention Grant, National Research Council of Canada Operating Grant No. A3881, National Science Foundation (U.S.A.) Operating Grant No. PCM74-13643-A02, and the Council of Forest Industries of British Columbia.     

Impact of Botanical Pesticides Derived from Melia azedarach and Azadirachta indica Plants on the Emission of Volatiles that Attract Parasitoids of the Diamondback Moth to Cabbage Plants

Herbivorous and carnivorous arthropods use chemical information from plants during foraging. Aqueous leaf extracts from the syringa tree Melia azedarach and commercial formulations from the neem tree Azadirachta indica, Neemix 4.5^®, were investigated for their impact on the flight response of two parasitoids, Cotesia plutellae and Diadromus collaris. Cotesia plutellae was attracted only to Plutella xylostella-infested cabbage plants in a wind tunnel after an oviposition experience. Female C. plutellae did not distinguish between P. xylostella-infested cabbage plants treated with neem and control P. xylostella-infested plants. However, females preferred infested cabbage plants that had been treated with syringa extract to control infested plants. Syringa extract on filter paper did not attract C. plutellae. This suggests that an interaction between the plant and the syringa extract enhances parasitoid attraction. Diadromus collaris was not attracted to cabbage plants in a wind tunnel and did not distinguish between caterpillar-damaged and undamaged cabbage plants. Headspace analysis revealed 49 compounds in both control cabbage plants and cabbage plants that had been treated with the syringa extract. Among these are alcohols, aldehydes, ketones, esters, terpenoids, sulfides, and an isothiocyanate. Cabbage plants that had been treated with the syringa extract emitted larger quantities of volatiles, and these increased quantities were not derived from the syringa extract. Therefore, the syringa extract seemed to induce the emission of cabbage volatiles. To our knowledge, this is the first example of a plant extract inducing the emission of plant volatiles in another plant. This interesting phenomenon likely explains the preference of C. plutellae parasitoids for cabbage plants that have been treated with syringa extracts.     

Volatile Stimuli Related to Feeding Activity of Nonprey Caterpillars, Spodoptera exigua, Affect Olfactory Response of the Predatory Mite Phytoseiulus persimilis

The effect of volatiles related to feeding activity of nonprey caterpillars, Spodoptera exigua, on the olfactory response of the predatory mites Phytoseiulus persimilis was examined in a Y-tube olfactometer. At a low caterpillar density (20 caterpillars on 10 Lima bean leaves), the predators were significantly more attracted to volatiles from infested leaves on which the caterpillars and their products were present or from infested leaves from which the caterpillars and their products had been removed when compared to volatiles from uninfested leaves. The predators, however, significantly avoided odors from 20 caterpillars and their products (mainly feces) removed from bean leaves. In contrast, at a higher caterpillar density (100 caterpillars on 10 Lima bean leaves), the predators avoided volatiles from caterpillar-infested bean leaves. Volatiles from infested leaves from which the caterpillars and their products had been removed were not preferred over volatiles from uninfested leaves. Volatiles from feces collected from 100 caterpillars were strongly avoided by the predators, while the behavior of the predatory mites was not affected by volatiles from 100 caterpillars removed from a plant. The data show that carnivorous arthropods may avoid nonprofitable herbivores. This avoidance seems to result from an interference of volatiles from herbivore products with the attraction to herbivore-induced plant volatiles.     

Induction and Systemic Release of Herbivore-Induced Plant Volatiles Mediating In-Flight Orientation of Aphidius ervi

In-flight orientation of the braconid Aphidius ervi in response to volatiles released from broad bean plants infested by the pea aphid, Acyrthosiphon pisum, was studied in a no-choice wind-tunnel bioassay. The role of aphid infestation level and duration, systemic production of volatiles by insect-free parts of the plant, and the specificity of aphid-induced volatiles on the flight behavior of the foraging female parasitoids were investigated. The upper insect-free part of a three-leaved broad bean plant, which was basally infested by a population of 40 A. pisum, released synomones detectable by A. ervi females after at least 48–72 hr of infestation, resulting in both significant increases in oriented flights and landings on the source compared with uninfested control plants. This suggests that volatiles involved in host-location by A. ervi are systemically released by broad bean plants either in response to circulation of aphid saliva, circulation of saliva-induced bioactive elicitors, or circulation of the synomones themselves. Air entrainment extracts of volatiles collected from a broad bean plant infested by the nonhost Aphis fabae or an uninfested broad bean plant elicited few oriented flights and landing responses by female parasitoids. These extracts were significantly less attractive than extracts collected from a broad bean plant infested by the host A. pisum, indicating the specificity of synomones elicited by different aphid species on the same plant species.     

Kairomones for the egg parasiteTrichogramma evanescens Westwood

In observation-cage experiments some new contact kairomones for the egg parasiteTrichogramma evanescens Westwood are demonstrated.T. evanescens females search significantly longer on cabbage leaves treated with the wing scales of two hosts,Pieris brassicae L. andP. rapae L. Further, egg washes ofP. brassicae containing an oviposition deterrent pheromone for the butterflies, were found to have a contact-kairomonal effect on the parasite.T. evanescens females search significantly longer on cabbage leaves sprayed with a methanol or water wash ofP. brassicae eggs than on leaves treated with the solvent only.Hymenoptera: Trichogrammatidae.Lepidoptera: Pieridae.     

A chemical basis for differential acceptance ofErysimum cheiranthoides by twoPieris species

Wormseed mustard,Erysimum cheiranthoides, is unacceptable as a host for the cabbage butterfly,Pieris rapae. However, it is preferred for oviposition byPieris napi oleracea in the greenhouse. Isolation and identification of the oviposition stimulants toP. napi oleracea were accomplished by C₁₈ open-column chromatography, TLC, ion-exchange chromatography, HPLC, UV, and NMR spectroscopy. Glucoiberin and glucocheirolin were identified as the most active stimulants. The extracted glucoiberin was as stimulatory as glucocheirolin, although its concentration in theErysimum plants was about 10 times lower than that of glucocheirolin. These glucosinolates were only weak stimulants toP. rapae. Furthermore,P. rapae was strongly deterred by the cardenolides, erysimoside and erychroside, fromE. cheiranthoides, andP. napi oleracea was less sensitive to these compounds. No other deterrent toP. napi oleracea was detected in this plant species. The results explain the differential acceptance ofE. cheiranthoides by these twoPieris species.     

Effects of molasses grass,Melinis minutiflora volatiles on the foraging behavior of the cereal stemborer parasitoid,Cotesia sesamiae

Olfactory responses of the cereal stemborer parasitoid Cotesia sesamiae to volatiles emitted by gramineous host and nonhost plants of the stemborers were studied in a Y-tube olfactometer. The host plants were maize (Zea mays) and sorghum ( Sorghum bicolor), while the nonhost plant was molasses grass (Melinis minutiflora). In single-choice tests, females of C. sesamiae chose volatiles from infested and uninfested host plants and molasses grass over volatiles from the control (soil). In dual-choice tests, the wasp preferred volatiles from infested host plants to those from uninfested host plants. There was no discrimination between molasses grass volatiles and those of uninfested maize, uninfested sorghum, or infested maize. The wasp preferred sorghum volatiles over maize. Combining uninfested maize or sorghum with molasses grass did not make volatiles from the combination more attractive as compared to only uninfested host plants. Infested maize alone was as attractive as when combined with molasses grass. Infested sorghum was preferred over its combination with molasses grass. Local growth conditions of the molasses grasses influenced attractiveness to the parasitoids. Volatiles from Thika molasses grass were attractive, while those from Mbita molasses grass were not. Growing the Thika molasses grass in Mbita rendered it unattractive and vice versa with the Mbita molasses grass. This is a case of the same genotype expressing different phenotypes due to environmental factors.     

Attraction of Dibrachys cavus (Hymenoptera: Pteromalidae) to its Host Frass Volatiles

The European grapevine moth, Lobesia botrana (Lepidoptera: Tortricidae), is a polyphagous insect able to develop on grapes and wild plants. We tested the hypothesis that the parasitoid Dibrachys cavus (Hymenoptera: Pteromalidae) uses the larval frass in its host search. A two-armed olfactometer was used to measure the attractiveness of L. botrana larvae, their silk, or their frass after larvae were fed on different host plants. Frass of three Lepidoptera (L. botrana, Eupoecillia ambiguella, Sphinx ligustri) and one Orthoptera (Chorthippus brunneus) was assayed, but only L. botrana was used to test an effect of the larval host plant (two grape cultivars and three other plant species) to D. cavus females. Larvae without frass did not attract D. cavus whatever their origin, but their frass was attractive at a dose of 2–3 days equivalent of larval frass production. The silk produced by a single larva (L. botrana) was not attractive to D. cavus. The parasitoid was most attracted to the odor of S. ligustri; the frass of L. botrana was more attractive than that of E. ambiguella, irrespective of the species on which D. cavus had been reared. There was no difference in attractiveness of frass collected from L. botrana raised on food containing different plants. Chemical extracts using five different polarity solvents (acetone, dichloromethane, hexane, methanol, and water) differed in attractiveness to D. cavus. Water and dichloromethane were the most attractive. This suggests that a complex volatile signal made from intermediate to polar volatiles may be involved in attraction. D. cavus used frass to discriminate between different potential host species. Our results revealed that the larval food of L. botrana did not modify frass attractiveness, but that the moth species did.     

Chemical constituents of an unacceptable crucifer,Erysimum cheiranthoides,deter feeding byPieris rapae

The wild cruciferErysimum cheiranthoides was found to contain extractable constituents that deterred feeding by larvae of the crucifer specialistPieris rapae when applied to cabbage leaf disks in both choice and nochoice bioassays. High-performance liquid chromatography was used to separate the extract into several fractions, two of which retained the feeding deterrent activity of the extract. UV-absorption spectra of the fractions suggested that one contained cardenolides similar or identical to those reported to deter oviposition byP. rapae onE. cheiranthoides. The other active fraction evidently contains a compound that deters larval feeding but not adult oviposition. The results suggest that the chemical defense ofE. cheiranthoides depends on two types of compounds acting on separate developmental stages of the insect.