Challenging the Preference-Performance Hypothesis in an above-belowground insect.

The relationship between female oviposition preference and offspring performance has been a question of special interest in the study of host plant selection by phytophagous insects. The Preference-Performance Hypothesis (PPH) is one of the main hypotheses proposed to explain this relationship, stating that females should preferentially lay eggs on plants providing the best larval development. The PPH has been extensively tested on aboveground insects but its application to species with belowground larvae is still mostly unknown. In this study, the PPH was quantitatively tested in an above-belowground insect, the cabbage root fly Delia radicum. Female oviposition preference and larval performance were estimated on three brassicaceous species (Brassica oleracea, Brassica rapa, and Sinapis alba) as well as between four cultivars of B. rapa and four cultivars of S. alba. Larval performance was estimated through their survival and through three life-history traits (LHT) of emerging adults. The PPH was supported at the intraspecific scale but only in B. rapa and for some, but not all, of the life-history traits. No support for the PPH was found in S. alba as well as at the interspecific scale. This study pleads for the integration of insects with both above- and belowground life stages in the preference-performance debate. Moreover, it raises the importance of measuring several variables to estimate larval performance and to test the PPH quantitatively, both at the plant intraspecific and interspecific scales, before drawing general conclusions.

Influence of the symbiont Wolbachia on life history traits of the cabbage root fly (Delia radicum).

Wolbachia is an endocellular bacteria infecting arthropods and nematodes and is only transmitted vertically by females via the cytoplasm of the egg. It is often a manipulator of host reproduction, causing cytoplasmic incompatibility, thelytokous parthenogenesis, feminization or male killing, which all increase the proportion of infected females in the population. However, Wolbachia can modify life history traits of the host without causing the above phenotypes and each species illustrates the variability of relationships between this remarkably versatile symbiont and its many hosts. We have measured maternal transmission and the impact of a natural Wolbachia infection in the cabbage root fly Delia radicum, a major agricultural pest. We used a population that is polymorphic for the infection to ensure similar genetic and microbiome backgrounds between groups. Maternal transmission of the infection was 100% in our sample. We found no evidence of cytoplasmic incompatibility, thelytokous parthenogenesis, feminization nor male killing. Wolbachia infection significantly reduced hatch rate in infected eggs (by 10%) but improved larvo-nymphal viability sufficiently so that infected eggs nevertheless yielded as many adults as uninfected ones, albeit with a 1.5% longer total development time. Starved and infected ovipositing females suffered significantly reduced viability (20% higher mortality during a 3-day oviposition period) than uninfected females, but mortality was not higher in starved virgin females nor in starved males, suggesting that the energetic cost of the infection is only revealed in extreme conditions. Wolbachia had no effect on egg hatch time or offspring size. The apparently 100% vertical transmission and the significant but mutually compensating effects found suggest that infection might be nearly benign in this host and might only drift slowly, which would explain why the infection rate has been stable in our laboratory (approximately 50% individuals infected) for at least 30 generations.

Characterizing volatiles and attractiveness of five brassicaceous plants with potential for a 'push-pull' strategy toward the cabbage root fly, Delia radicum.

Volatile Organic Compounds (VOCs) released by plants are involved in various orientation processes of herbivorous insects and consequently play a crucial role in their reproductive success. In the context of developing new strategies for crop protection, several studies have previously demonstrated the possibility to limit insect density on crops using either host or non-host plants that release attractive or repellent VOCs, respectively. The cabbage root fly, Delia radicum, is an important pest of brassicaceous crops for which control methods have to be implemented. Several studies have shown that plant odors influence cabbage root fly behavior, but only few VOCs have been identified so far. The present study aimed at selecting both plants and olfactory stimuli that could be used in the development of a "push-pull" strategy against the cabbage root fly. Olfactometer results revealed that plants belonging to the same family, even to the same species, may exhibit different levels of attractiveness toward D. radicum. Plants that were found attractive in behavioral observations were characterized by high release rates of distinct terpenes, such as linalool, ß-caryophyllene, humulene, and α-farnesene. This study represents a first step to identify both attractive plants of agronomic interest, and additional volatiles that could be used in the context of trap crops to protect broccoli fields against the cabbage root fly.

Interactions Between a Belowground Herbivore and Primary and Secondary Root Metabolites in Wild Cabbage.

Plants are attacked by both above- and belowground herbivores. Toxic secondary compounds are part of the chemical defense arsenal of plants against a range of antagonists, and are subject to genetic variation. Plants also produce primary metabolites (amino acids, nutrients, sugars) that function as essential compounds for growth and survival. Wild cabbage populations growing on the Dorset coast of the UK exhibit genetically different chemical defense profiles, even though they are located within a few kilometers of each other. As in other Brassicaceae, the defensive chemicals in wild cabbages constitute, among others, secondary metabolites called glucosinolates. Here, we used five Dorset populations of wild cabbage to study the effect of belowground herbivory by the cabbage root fly on primary and secondary chemistry, and whether differences in chemistry affected the performance of the belowground herbivore. There were significant differences in total root concentrations and chemical profiles of glucosinolates, amino acids, and sugars among the five wild cabbage populations. Glucosinolate concentrations not only differed among the populations, but also were affected by root fly herbivory. Amino acid and sugar concentrations also differed among the populations, but were not affected by root fly herbivory. Overall, population-related differences in plant chemistry were more pronounced for the glucosinolates than for amino acids and sugars. The performance of the root herbivore did not differ among the populations tested. Survival of the root fly was low (<40%), suggesting that other belowground factors may override potential differences in effects related to primary and secondary chemistry.

Manipulating feeding stimulation to protect crops against insect pests?

Enhancing natural mechanisms of plant defense against herbivores is one of the possible strategies to protect cultivated species against insect pests. Host plant feeding stimulation, which results from phagostimulant and phagodeterrent effects of both primary and secondary metabolites, could play a key role in levels of damage caused to crop plants. We tested this hypothesis by comparing the feeding intensity of the pollen beetle Meligethes aeneus on six oilseed rape (Brassica napus) genotypes in a feeding experiment, and by assessing the content of possible phagostimulant and phagodeterrent compounds in tissues targeted by the insect (flower buds). For this purpose, several dozens of primary and secondary metabolites were quantified by a set of chromatographic techniques. Intergenotypic variability was found both in the feeding experiment and in the metabolic profile of plant tissues. Biochemical composition of the perianth was in particular highly correlated with insect damage. Only a few compounds explained this correlation, among which was sucrose, known to be highly phagostimulating. Further testing is needed to validate the suggested impact of the specific compounds we have identified. Nevertheless, our results open the way for a crop protection strategy based on artificial selection of key determinants of insect feeding stimulation.

Parasitoid-specific induction of plant responses to parasitized herbivores affects colonization by subsequent herbivores.

Plants are exposed to a suite of herbivorous attackers that often arrive sequentially. Herbivory affects interactions between the host plants and subsequently attacking herbivores. Moreover, plants may respond to herbivory by emitting volatile organic compounds (VOCs) that attract carnivorous natural enemies of the herbivores. However, information borne by VOCs is ubiquitous and may attract carnivores, such as parasitoids, that differ in their effectiveness at releasing the plant from its herbivorous attackers. Furthermore, the development of parasitoids within their herbivorous hosts, attacking a given host plant, may influence the elicitation of defensive reactions in the host plant. This may, in turn, affect the behavior of subsequent herbivores attacking the host plant. Here, we show that the species identity of a parasitoid had a more significant effect on defense responses of Brassica oleracea plants than the species identity of the herbivorous hosts of the parasitoids. Consequently, B. oleracea plants that were damaged by caterpillars (Pieris spp.) parasitized by different parasitoid species varied in the degree to which diamondback moths (Plutella xylostella) selected the plants for oviposition. Attracting parasitoids in general benefitted the plants by reducing diamondback moth colonization. However, the species of parasitoid that parasitized the herbivore significantly affected the magnitude of this benefit by its species-specific effect on herbivore-plant interactions mediated by caterpillar regurgitant. Our findings show that information-mediated indirect defense may lead to unpredictable consequences for plants when considering trait-mediated effects of parasitized caterpillars on the host plant and their consequences because of community-wide responses to induced plants.

Understanding crop colonization of oilseed rape crops by the cabbage stem flea beetle (Psylliodes chrysocephala L. (Coleoptera: Chrysomelidae)).

BACKGROUND: Development of accurate pest monitoring systems is essential for the establishment of integrated pest management strategies. Information about the pest behavior during the colonization process, as well as the sex and reproductive status of the colonizing population often are lacking and hinder their development. The cabbage stem flea beetle (CSFB, Psylliodes chrysocephala) can cause the complete destruction of oilseed rape crops (OSR, Brassica napus). In the present study, the colonization process of OSR fields by the CSFB was studied. RESULTS: More individuals were caught on the outward facing side of the traps than the side of the trap facing towards the crop at the field border and catches were higher on the trapping units at the center of the field than at its border, suggesting that more beetles were entering than leaving the crop. Catches were higher on lower traps placed near to the crop than on those positioned further from the ground and also were higher during the day than late afternoon and night. The sex-ratio of individuals caught was skewed towards males and sexual maturity was acquired for females during the experiment. Integration of sampling data with local meteorological data showed that the catches correlated mostly with air temperature and relative humidity. CONCLUSION: This study provides new information about the dispersion of the CSFB in OSR fields during the colonization process, and highlights correlations between local meteorological factors and activity of the CSFB, and represent a new step towards implementing monitoring strategies against this pest. © 2023 The Authors. Pest Management Science published by John Wiley & Sons Ltd on behalf of Society of Chemical Industry.

A novel indirect defence in Brassicaceae: structure and function of extrafloral nectaries in Brassica juncea.

While nectaries are commonly found in flowers, some plants also form extrafloral nectaries on stems or leaves. For the first time in the family Brassicaceae, here we report extrafloral nectaries in Brassica juncea. The extrafloral nectar (EFN) was secreted from previously amorphic sites on stems, flowering stalks and leaf axils from the onset of flowering until silique formation. Transverse sections at the point of nectar secretion revealed a pocket-like structure whose opening was surrounded by modified stomatal guard cells. The EFN droplets were viscous and up to 50% of the total weight was sugars, 97% of which was sucrose in the five varieties of B. juncea examined. Threonine, glutamine, arginine and glutamate were the most abundant amino acids. EFN droplets also contained glucosinolates, mainly gluconapin and sinigrin. Nectar secretion was increased when the plants were damaged by chewing above- and belowground herbivores and sap-sucking aphids. Parasitoids of each herbivore species were tested for their preference, of which three parasitoids preferred EFN and sucrose solutions over water. Moreover, the survival and fecundity of parasitoids were positively affected by feeding on EFN. We conclude that EFN production in B. juncea may contribute to the indirect defence of this plant species.

Age-specific allocation of glucosinolates within plant reproductive tissues.

The Optimal Defense Theory (ODT) predicts that the distribution of defenses within a plant should mirror the value and vulnerability of each tissue. Although the ODT has received much experimental support, very few studies have examined defense allocation among reproductive tissues and none assessed simultaneously how these defenses evolve with age. We quantified glucosinolates in perianths, anthers and pistils at different bud maturity stages (i.e., intermediate flower buds, old flower buds and flowers) of undamaged and mechanically damaged plants of an annual brassicaceous species. The youngest leaf was used as a reference for vegetative organs, since it is predicted to be one of the most defended. In line with ODT predictions, reproductive tissues were more defended than vegetative tissues constitutively, and within the former, pistils and anthers more defended than perianths. No change in the overall defense level was found between bud maturity stages, but a significant temporal shift was observed between pistils and anthers. Contrary to ODT predictions, mechanical damage did not induce systemic defenses in leaves but only in pistils. Our results show that defense allocation in plant reproductive tissues occurs at fine spatial and temporal scales, extending the application framework of the ODT. They also demonstrate interactions between space and time in fine-scale defense allocation.

Plasticity in Chemical Host Plant Recognition in Herbivorous Insects and Its Implication for Pest Control.

Chemical communication is very important in herbivorous insects, with many species being important agricultural pests. They often use olfactory cues to find their host plants at a distance and evaluate their suitability upon contact with non-volatile cues. Responses to such cues are modulated through interactions between various stimuli of biotic and abiotic origin. In addition, the response to the same stimulus can vary as a function of, for example, previous experience, age, mating state, sex, and morph. Here we summarize recent advances in the understanding of plant localization and recognition in herbivorous insects with a focus on the interplay between long- and short-range signals in a complex environment. We then describe recent findings illustrating different types of plasticity in insect plant choice behavior and the underlying neuronal mechanisms at different levels of the chemosensory pathway. In the context of strong efforts to replace synthetic insecticides with alternative pest control methods, understanding combined effects between long- and close-range chemical cues in herbivore-plant interactions and their complex environment in host choice are crucial to develop effective plant protection methods. Furthermore, plasticity of behavioral and neuronal responses to chemical cues needs to be taken into account to develop effective sustainable pest insect control through behavioral manipulation.

A protocol for increased throughput phenotyping of plant resistance to the pollen beetle.

BACKGROUND: Improving crop resistance to insect herbivores is a major research objective in breeding programs. Although genomic technologies have increased the speed at which large populations can be genotyped, breeding programs still suffer from phenotyping constraints. The pollen beetle (Brassicogethes aeneus) is a major pest of oilseed rape for which no resistant cultivar is available to date, but previous studies have highlighted the potential of white mustard as a source of resistance and introgression of this resistance appears to be a promising strategy. Here we present a phenotyping protocol allowing mid-throughput (i.e., increased throughput compared to current methods) acquisition of resistance data, which could then be used for genetic mapping of QTLs. RESULTS: Contrasted white mustard genotypes were selected from an initial field screening and then evaluated for their resistance under controlled conditions using a standard phenotyping method on entire plants. We then upgraded this protocol for mid-throughput phenotyping, by testing two alternative methods. We found that phenotyping on detached buds did not provide the same resistance contrasts as observed with the standard protocol, in contrast to the phenotyping protocol with miniaturized plants. This protocol was then tested on a large panel composed of hundreds of plants. A significant variation in resistance among genotypes was observed, which validates the large-scale application of this new phenotyping protocol. CONCLUSION: The combination of this mid-throughput phenotyping protocol and white mustard as a source of resistance against the pollen beetle offers a promising avenue for breeding programs aiming to improve oilseed rape resistance. © 2022 The Authors. Pest Management Science published by John Wiley & Sons Ltd on behalf of Society of Chemical Industry.

A high-quality functional genome assembly of Delia radicum L. (Diptera: Anthomyiidae) annotated from egg to adult.

Belowground herbivores are overseen and underestimated, even though they can cause significant economic losses in agriculture. The cabbage root fly Delia radicum (Anthomyiidae) is a common pest in Brassica species, including agriculturally important crops, such as oilseed rape. The damage is caused by the larvae, which feed specifically on the taproots of Brassica plants until they pupate. The adults are aboveground-living generalists feeding on pollen and nectar. Female flies are attracted by chemical cues in Brassica plants for oviposition. An assembled and annotated genome can elucidate which genetic mechanisms underlie the adaptation of D. radicum to its host plants and their specific chemical defences, in particular isothiocyanates. Therefore, we assembled, annotated and analysed the D. radicum genome using a combination of different next-generation sequencing and bioinformatic approaches. We assembled a chromosome-level D. radicum genome using PacBio and Hi-C Illumina sequence data. Combining Canu and 3D-DNA genome assembler, we constructed a 1.3 Gbp genome with an N50 of 242 Mbp and 6 pseudo-chromosomes. To annotate the assembled D. radicum genome, we combined homology-, transcriptome- and ab initio-prediction approaches. In total, we annotated 13,618 genes that were predicted by at least two approaches. We analysed egg, larval, pupal and adult transcriptomes in relation to life-stage specific molecular functions. This high-quality annotated genome of D. radicum is a first step to understanding the genetic mechanisms underlying host plant adaptation. As such, it will be an important resource to find novel and sustainable approaches to reduce crop losses to these pests.

Differences in volatile profiles of turnip plants subjected to single and dual herbivory above- and belowground.

Plants attacked by herbivorous insects emit volatile organic compounds that are used by natural enemies to locate their host or prey. The composition of the blend is often complex and specific. It may vary qualitatively and quantitatively according to plant and herbivore species, thus providing specific information for carnivorous arthropods. Most studies have focused on simple interactions that involve one species per trophic level, and typically have investigated the aboveground parts of plants. These investigations need to be extended to more complex networks that involve multiple herbivory above- and belowground. A previous study examined whether the presence of the leaf herbivore Pieris brassicae on turnip plants (Brassica rapa subsp. rapa) influences the response of Trybliographa rapae, a specialist parasitoid of the root feeder Delia radicum. It showed that the parasitoid was not attracted by volatiles emitted by plants under simultaneous attack. Here, we analyzed differences in the herbivore induced plant volatile (HIPV) mixtures that emanate from such infested plants by using Orthogonal Partial Least Squares-Discriminant Analysis (OPLS-DA). This multivariate model focuses on the differences between odor blends, and highlights the relative importance of each compound in an HIPV blend. Dual infestation resulted in several HIPVs that were present in both isolated infestation types. However, HIPVs collected from simultaneously infested plants were not the simple combination of volatiles from isolated forms of above- and belowground herbivory. Only a few specific compounds characterized the odor blend of each type of damaged plant. Indeed, some compounds were specifically induced by root herbivory (4-methyltridecane and salicylaldehyde) or shoot herbivory (methylsalicylate), whereas hexylacetate, a green leaf volatile, was specifically induced after dual herbivory. It remains to be determined whether or not these minor quantitative variations, within the background of more commonly induced odors, are involved in the reduced attraction of the root feeder's parasitoid. The mechanisms involved in the specific modification of the odor blends emitted by dual infested turnip plants are discussed in the light of interferences between biosynthetic pathways linked to plant responses to shoot or root herbivory.

Influential Insider: Wolbachia, an Intracellular Symbiont, Manipulates Bacterial Diversity in Its Insect Host.

Facultative intracellular symbionts like the α-proteobacteria Wolbachia influence their insect host phenotype but little is known about how much they affect their host microbiota. Here, we quantified the impact of Wolbachia infection on the bacterial community of the cabbage root fly Delia radicum by comparing the microbiota of Wolbachia-free and infected adult flies of both sexes. We used high-throughput DNA sequencing (Illumina MiSeq, 16S rRNA, V5-V7 region) and performed a community and a network analysis. In both sexes, Wolbachia infection significantly decreased the diversity of D. radicum bacterial communities and modified their structure and composition by reducing abundance in some taxa but increasing it in others. Infection by Wolbachia was negatively correlated to 8 bacteria genera (Erwinia was the most impacted), and positively correlated to Providencia and Serratia. We suggest that Wolbachia might antagonize Erwinia for being entomopathogenic (and potentially intracellular), but would favor Providencia and Serratia because they might protect the host against chemical plant defenses. Although they might seem prisoners in a cell, endocellular symbionts can impact the whole microbiota of their host, hence its extended phenotype, which provides them with a way to interact with the outside world.

Kin recognition loss following anesthesia in beetle larvae (Aleochara bilineata, Coleoptera, Staphylinidae).

Kin recognition is a complex cognitive process, where an individual should detect a phenotypic cue and compare it to an internal template, which might be genetically determined (i.e., innate or acquired). Kin recognition mechanisms will depend on whether previous encounters with kin are possible or not to form the individual internal template. When relatives have never met before, kin recognition is supposed to rely on recognition alleles (which allows the innate recognition of relatives bearing them), or on self-referent phenotype matching (the individual has formed a template using its own phenotype and recognizes as kin individuals which match it closely enough). Although self-referent phenotype matching is in theory the more likely explanation, it has not been possible so far to exclude experimentally the recognition alleles' hypothesis. Here, we report that kin recognition in the solitary parasitoid larvae of Aleochara bilineata (Coleoptera; Staphylinidae) is suppressed following carbon dioxide anesthesia or chill-coma, both treatments known to cause a temporary amnesia. Treated larvae superparasitize indifferently hosts parasitized either by siblings or by non-kin larvae, while untreated larvae avoid hosts occupied by siblings. The two types of anesthesia thus suppress kin recognition, but their global effect on larvae is different. Chill-coma suppresses the ability to distinguish parasitized from unparasitized hosts and reduces parasitism rate, suggesting an aspecific impairment of sensory receptors or cognition. However, carbon dioxide narcosis only impairs kin recognition, strongly suggesting that an intact memory is necessary for kin recognition to take place. Although this study does not address the recognition alleles' hypothesis per se, our results strongly support a self-referent phenotype matching mechanism. On the whole, kin recognition in A. bilineata larvae is effective through short-term memory, because it is affected by amnesic treatments.

Long-lasting effects of antibiotics on bacterial communities of adult flies.

Insect symbionts benefit their host and their study requires large spectrum antibiotic use like tetracycline to weaken or suppress symbiotic communities. While antibiotics have a negative impact on insect fitness, little is known about antibiotic effects on insect microbial communities and how long they last. We characterized the bacterial communities of adult cabbage root fly Delia radicum in a Wolbachia-free population and evaluated the effect of tetracycline treatment on these communities over several generations. Three D. radicum generations were used: the first- and second-generation flies either ingested tetracycline or not, while the third-generation flies were untreated but differed with their parents and/or grandparents that had or had not been treated. Fly bacterial communities were sequenced using a 16S rRNA gene. Tetracycline decreased fly bacterial diversity and induced modifications in both bacterial abundance and relative frequencies, still visible on untreated offspring whose parents and/or grandparents had been treated, therefore demonstrating long-lasting transgenerational effects on animal microbiomes after antibiotic treatment. Flies with an antibiotic history shared bacterial genera, potentially tetracycline resistant and heritable. Next, the transmission should be investigated by comparing several insect development stages and plant compartments to assess vertical and horizontal transmissions of D. radicum bacterial communities.

The Potential for Decision Support Tools to Improve the Management of Root-Feeding Fly Pests of Vegetables in Western Europe.

Several important vegetable crops grown outdoors in temperate climates in Europe can be damaged by the root-feeding larvae of Diptera (Delia radicum, Delia floralis, Chamaepsila rosae, Delia platura, Delia florilega, Delia antiqua). Knowledge of pest insect phenology is a key component of any Integrated Pest Management (IPM) strategy, and this review considers the methods used to monitor and forecast the occurrence of root-feeding flies as a basis for decision-making by growers and the ways that such information can be applied. It has highlighted some current management approaches where such information is very useful for decision support, for example, the management of C. rosae with insecticidal sprays and the management of all of these pests using crop covers. There are other approaches, particularly those that need to be applied at sowing or transplanting, where knowledge of pest phenology and abundance is less necessary. Going forward, it is likely that the number of insecticidal control options available to European vegetable growers will diminish and they will need to move from a strategy which often involves using a single 'silver bullet' to a combination of approaches/tools with partial effects (applied within an IPM framework). For the less-effective, combined methods, accurate information about pest phenology and abundance and reliable decision support are likely to be extremely important.

Oviposition Preference of the Cabbage Root Fly towards Some Chinese Cabbage Cultivars: A Search for Future Trap Crop Candidates.

The development of integrated pest management strategies becomes more and more pressing in view of potential harmful effects of synthetic pesticides on the environment and human health. A promising alternative strategy against Delia radicum is the use of trap crops. Chinese cabbage (Brassica rapa subsp. pekinensis and subsp. chinensis) is a highly sensitive Brassicaceae species previously identified as a good candidate to attract the cabbage root fly away from other crops. Here, we carried out multi-choice experiments both in the laboratory and in field conditions to measure the oviposition susceptibilities of different subspecies and cultivars of Chinese cabbages as compared to a broccoli reference. We found large differences among subspecies and cultivars of the Chinese cabbage, which received three to eleven times more eggs than the broccoli reference in field conditions. In laboratory conditions, the chinensis subspecies did not receive more eggs than the broccoli reference. We conclude that D. radicum largely prefers to lay eggs on the pekinensis subspecies of Chinese cabbage compared to the chinensis subspecies or broccoli. Some pekinensis cultivars, which received over ten times more eggs than broccoli in the field, appear especially promising candidates to further develop trap crop strategies against the cabbage root fly.

Early Olfactory Environment Influences Antennal Sensitivity and Choice of the Host-Plant Complex in a Parasitoid Wasp.

Early experience of olfactory stimuli associated with their host-plant complex (HPC) is an important driver of parasitoid foraging choices, notably leading to host fidelity. Mechanisms involved, such as peripheral or central modulation, and the impact of a complex olfactory environment are unknown. Using olfactometer assays, we compared HPC preference of Aphidius ervi Haliday (Hymenoptera:Braconidae) females originating from two different HPCs, either with the other HPC in close vicinity (complex environment) or without (simple environment). We also investigated antennal responses to volatiles differentially emitted by the two respective HPCs. In a simple environment, HPC of origin had an influence on olfactory choice, but the preferences observed were asymmetric according to parasitoid origin. Electroantennographic recordings revealed significant sensitivity differences for some of the tested individual volatiles, which are emitted differentially by the two HPCs. Besides, presence of an alternative HPC during early stages modified subsequent parasitoid preferences. We discuss how increased olfactory complexity could influence parasitoid host foraging and biological control in diversified cropping systems.

Kin recognition in Aleochara bilineata could support the kinship theory of genomic imprinting.

Genomic imprinting corresponds to the differential expression of a gene according to its paternal or maternal origin. The kinship theory of genomic imprinting proposes that maternally or paternally inherited genes may be in conflict over their effects on kin differently related along the paternal or maternal line. Most examples supporting the kinship theory of imprinting deal with competition between offspring for maternal resources. However, genomic imprinting may also explain differential behavioral expression toward kin whenever sibs are more related to each other via one parental sex than the other. Unfortunately, nothing is currently known about imprinting associated with a behavioral phenotype in insects. Here we report the first evidence of such a maternally imprinted behavior. We show that the solitary parasitoid larvae of Aleochara bilineata Gyll (Coleoptera; Staphylinidae), which avoid superparasitizing their full sibs, also avoid their cousins when they are related to them through their father, but not when they are related to them through their mother. A genetic kin recognition mechanism is proposed to explain this result and we conclude that genomic imprinting could control the avoidance of kin superparasitism in this species and have a profound influence on decision-making processes.