Responses of green lacewings to semiochemicals: species- and sex-specificity (Neuroptera: Chrysopidae)

ABSTRACT Green lacewings (Chrysopidae) are important predators in agroecosystems. Frequently there are studies which treat these beneficial insects at family level as a whole. This approach, despite its practical advantages, may, however, include many species with markedly different life history traits. As green lacewings are also rather diverse in their chemical ecology, treating several different species within one unit may lead to confusion and confounding effects of natural phenomena in research. Besides interspecific differences, sex-specificity is another important factor to consider in respect of insect chemical ecology. This is especially important for green lacewings, in particular the attraction of females, since oviposition is crucial in their application as biological control agents. The aim of our paper is to provide a brief insight into the diversity of the chemical ecology of green lacewings with a special emphasis on the species- and sex-specific differences in their responses to semiochemicals.

Attraction of Chrysotropia ciliata (Neuroptera, Chrysopidae) Males to P-Anisaldehyde, a Compound with Presumed Pheromone Function.

In a field-trapping experiment with plant volatiles, we observed notably high attraction of green lacewing (Chrysotropia ciliata) males to the compound p-anisaldehyde. Based on this finding, we initiated the present study to elucidate this phenomenon and to investigate the chemical ecology of C. ciliata. Scanning electron microscopy revealed elliptical glands abundantly distributed on the 2nd to 6th abdominal sternites of C. ciliata males, whereas females of the species completely lacked such glands. No p-anisaldehyde was found in extractions of body parts of C. ciliata. Methyl p-anisate and p-methoxybenzoic acid were identified exclusively in the extract from abdominal segments 2-8 of males. Field-trapping experiments revealed no attraction of C. ciliata to either methyl p-anisate or p-methoxybenzoic acid. In contrast, males showed marked attraction to p-anisaldehyde in the field and antennae showed strong responses to this compound. Headspace collections in the field from living insects in their natural environment and during their main daily activity period indicated that p-anisaldehyde was emitted exclusively by C. ciliata males. Our overall results suggest that p-anisaldehyde might serve as a male-produced pheromone that attracts conspecific C. ciliata males. Here, we discuss hypotheses regarding possible mechanisms involved in regulation of p-anisaldehyde production, including involvement of the compounds methyl p-anisate and p-methoxybenzoic acid, and the potential ecological function of p-anisaldehyde in C. ciliata.

New perspectives for simultaneous attraction of Chrysoperla and Chrysopa lacewing species for enhanced biological control (Neuroptera: Chrysopidae).

Green lacewings (Chrysopidae) are important predators of many soft-bodied pest insects, for instance aphids. Previous studies reported attraction of Chrysoperla carnea species-complex to a ternary floral bait. The larvae of these lacewings are important generalist predators in agroecosystems, however adults are non-predatory, they feed on pollen, nectar or honeydew. Squalene, a plant originated compound was previously reported to be attractive to the nearctic Chrysopa nigricornis. In the current study squalene was tested alone and in combination with the ternary bait in field experiments in Hungary. In our experiments, traps baited with squalene attracted predatory males of Chrysopa formosa. Traps baited with squalene and the ternary floral bait attracted adults of both C. formosa and C. carnea complex lacewings. To our knowledge this is the first report of a bait combination attractive to both Chrysoperla and Chrysopa species. This finding is of special interest considering the remarkably different feeding habits of the adults of these lacewings. Potential perspectives in biological control are discussed.

Conspecific and Heterogeneric Lacewings Respond to (Z)-4-Tridecene Identified from Chrysopa formosa (Neuroptera: Chrysopidae).

Green lacewings (Chrysopidae) are predators of soft-bodied pest insects and are among the most important biological control agents in crop protection. Chrysopa spp. are of special importance since, unlike most green lacewing species, adults are also predatory. The current study was undertaken in search of Chrysopa formosa compounds with semiochemical activity. Using coupled gas chromatography-electroantennography (GC-EAG), head and thorax extracts of C. formosa elicited EAG responses to a compound subsequently identified by coupled GC/mass spectrometry, microchemistry, chemical synthesis and GC peak enhancement as (Z)-4-tridecene. In field experiments, this compound decreased attraction of adult C. formosa to (1R,4aS,7S,7aR)-nepetalactol and that of Chrysoperla carnea species-complex to a ternary floral lure, with the inhibitory effect found to be dose-dependent. Our results suggest that (Z)-4-tridecene may serve as a general warning signal among multiple green lacewing species. Perspectives for potential practical applications are discussed.

Aphid Sex Pheromone Compounds Interfere with Attraction of Common Green Lacewings to Floral Bait.

Common green lacewings (Chrysoperla carnea complex) form a group of generalist predators important for biological control. Several reports show attraction of these insects to plant volatiles, and a highly attractive ternary compound floral bait has been developed. With aphids being a preferred prey of larvae, one might expect these lacewings to be attracted to aphid semiochemicals, for instance, to aphid sex pheromones, as found for several other green lacewing species. However, in a previous study, we found that traps containing aphid sex pheromone compounds (1R,4aS,7S,7aR)-nepetalactol (NEPOH), (4aS,7S,7aR)-nepetalactone (NEPONE), and a ternary floral bait attracted fewer individuals than those containing the ternary floral bait alone. In the present study, possible causes for this effect of NEPOH and NEPONE on trap capture were studied. We established that C. carnea complex catches in traps with a ternary floral lure were not influenced by the presence of Chrysopa formosa individuals in traps (attracted by NEPOH and NEPONE) or by synthetic skatole (a characteristic component of Chrysopa defense secretion). A direct negative effect of NEPOH and NEPONE on attraction of C. carnea complex was found, suggesting active avoidance of these aphid sex pheromone components. This finding is surprising as the larvae of these lacewings prey preferentially on aphids. Possible mechanisms underlying this phenomenon are discussed.

Attraction of Chrysoperla carnea complex and Chrysopa spp. lacewings (Neuroptera: Chrysopidae) to aphid sex pheromone components and a synthetic blend of floral compounds in Hungary.

BACKGROUND: The deployment of synthetic attractants for the manipulation of lacewing populations as aphid predators is currently used in integrated pest management. This study investigates a synthetic bait comprising floral compounds previously found to attract the Chrysoperla carnea complex, and, for the first time, the aphid sex pheromone components (1R,4aS,7S,7aR)-nepetalactol and (4aS,7S,7aR)-nepetalactone, in field experiments in Hungary, for their ability to manipulate lacewing populations. RESULTS: The synthetic floral bait attracted both sexes of the Chrysoperla carnea complex, and Chrysopa formosa Brauer showed minimal attraction. The aphid sex pheromone compounds alone attracted males of C. formosa and C. pallens (Rambur). When the two baits were combined, Chrysopa catches were similar to those with aphid sex pheromone baits alone, but carnea complex catches decreased significantly (by 85-88%). CONCLUSION: As the floral bait alone attracted both sexes of the carnea complex, it showed potential to manipulate the location of larval density via altering the site of oviposition. Aphid sex pheromone compounds alone attracted predatory males of Chrysopa spp. and can potentially be used to enhance biological control of aphids. For the carnea complex, however, a combination of both baits is not advantageous because of the decrease in adults attracted. Assumptions of intraguild avoidance underlying this phenomenon are discussed.

Optimization of a phenylacetaldehyde-based attractant for common green lacewings (Chrysoperla carnea s.l.).

In field trapping tests, the catch of Chrysoperla carnea sensu lato (Neuroptera: Chrysopidae) increased when acetic acid was added to lures with phenylacetaldehyde. The addition of methyl salicylate to the binary mixture of phenylacetaldehyde plus acetic acid increased catches even further. The ternary blend proved to be more attractive than beta-caryophyllene, 2-phenylethanol, or 3-methyl eugenol (compounds previously described as attractants for chrysopids) on their own, and no influence on catches was recorded when these compounds were added as fourth components to the ternary blend. There were minimal changes in activity when (E)-cinnamaldehyde or methyl anthranylate (both evoking large responses from female or male antennae of C. carnea in this study) were added, although both compounds showed significant attraction on their own when compared to unbaited traps. In subtractive field bioassays with the ternary mixture, it appeared that the presence of either phenylacetaldehyde or methyl salicylate was important, whereas acetic acid was less so in the ternary mixture. The ternary blend attracted both female and male lacewings at sites in southern, central, and northern Europe. Possible applications of a synthetic attractant for lacewings are discussed.

Spider web survey or whole plant visual sampling? Impact assessment of Bt corn on non-target predatory insects with two concurrent methods.

Impact assessment studies rely on relevant sampling methods. Correct evaluation of methods can be done by their concurrent use in the same sampling site and period. Collecting webs of Theridion impressum L. Koch (Araneae: Theridiidae) may serve as an arthropod sampling method: empty cuticles of prey items remain attached to the back side of the leaves with adhesive silk. Our study aimed to compare the applicability of the two methods concurrently (spider web survey and whole plant visual sampling) in a risk assessment study. The spider web survey recorded more predatory insect families than the whole plant visual sampling. Both methods were able to detect significant differences in the quantity of predatory insects in Bt vs. isogenic plots, but not in the same taxa (Nabidae: spider-web, 2001, Bt > Iso; Coccinellidae: plant sampling, 2001, Iso > Bt; Welsh-test, P < 0.05); thus, they could not confirm each other. The lack of confirmation can be explained by differences in the sensitivity and selectivity of the two methods. A web survey of T. impressum has the practical advantage that although we concentrate only on the one species during field sampling, we gain additional information on a wide range of foliage-dwelling arthropods. Due to several biological uncertainties, interpretation and explanation of the results remain problematic. Thus, additional research--based on in-situ observation and recording of T. impressum-prey interactions--is necessary before we could propose web survey method as a complementary tool in ecological impact assessment.