Competitive interactions in insect parasitoids: effects of microbial symbionts across tritrophic levels.

Competition for hosts is a common ecological interaction in insect parasitoids. In the recent years, it has become increasingly evident that microorganisms can act as 'hidden players' in parasitoid ecology. In this review, we propose that parasitoid competition should take into consideration the microbial influence. In particular, we take a tritrophic perspective and discuss how parasitoid competition can be modulated by microorganisms associated with the parasitoids, their herbivore hosts, or the plants attacked by the herbivores. Although research is still in its infancy, recent studies have shown that microbial symbionts can modulate the contest outcome. The emerging pattern is that microorganisms not only affect the competitive traits of parasitoids but also the fighting arena (i.e. the herbivore host and its food plant), in which competition takes place. We have also identified important gaps in the literature that should be addressed in future studies to advance our understanding about parasitoid competition.

Nectar-Inhabiting Bacteria Affect Olfactory Responses of an Insect Parasitoid by Altering Nectar Odors.

Floral nectar is ubiquitously colonized by a variety of microorganisms among which yeasts and bacteria are the most common. Microorganisms inhabiting floral nectar can alter several nectar traits, including nectar odor by producing microbial volatile organic compounds (mVOCs). Evidence showing that mVOCs can affect the foraging behavior of insect pollinators is increasing in the literature, whereas the role of mVOCs in altering the foraging behavior of third-trophic level organisms such as insect parasitoids is largely overlooked. Parasitoids are frequent visitors of flowers and are well known to feed on nectar. In this study, we isolated bacteria inhabiting floral nectar of buckwheat, Fagopyrum esculentum (Polygonales: Polygonaceae), to test the hypothesis that nectar bacteria affect the foraging behavior of the egg parasitoid Trissolcus basalis (Hymenoptera: Scelionidae) via changes in odors of nectar. In behavioral assays, we found that T. basalis wasps are attracted toward nectar fermented by 4 out of the 14 bacterial strains isolated, which belong to Staphylococcus epidermidis, Terrabacillus saccharophilus (both Firmicutes), Pantoea sp. (Proteobacteria), and Curtobacterium sp. (Actinobacteria). Results of chemical investigations revealed significant differences in the volatile blend composition of nectars fermented by the bacterial isolates. Our results indicate that nectar-inhabiting bacteria play an important role in the interactions between flowering plants and foraging parasitoids. These results are also relevant from an applied perspective as flowering resources, such as buckwheat, are largely used in agriculture to promote conservation biological control of insect pests.

Chemical Ecology of Floral Resources in Conservation Biological Control.

Conservation biological control aims to enhance populations of natural enemies of insect pests in crop habitats, typically by intentional provision of flowering plants as food resources. Ideally, these flowering plants should be inherently attractive to natural enemies to ensure that they are frequently visited. We review the chemical ecology of floral resources in a conservation biological control context, with a focus on insect parasitoids. We highlight the role of floral volatiles as semiochemicals that attract parasitoids to the food resources. The discovery that nectar-inhabiting microbes can be hidden players in mediating parasitoid responses to flowering plants has highlighted the complexity of the interactions between plants and parasitoids. Furthermore, because food webs in agroecosystems do not generally stop at the third trophic level, we also consider responses of hyperparasitoids to floral resources. We thus provide an overview of floral compounds as semiochemicals from a multitrophic perspective, and we focus on the remaining questions that need to be addressed to move the field forward.

Diversity and composition of the microbiome associated with eggs of the Southern green stinkbug, Nezara viridula (Hemiptera: Pentatomidae).

Although microbial communities of insects from larval to adult stage have been increasingly investigated in recent years, little is still known about the diversity and composition of egg-associated microbiomes. In this study, we used high-throughput amplicon sequencing and quantitative PCR to get a better understanding of the microbiome of insect eggs and how they are established using the Southern green stinkbug Nezara viridula (L.) (Hemiptera: Pentatomidae) as a study object. First, to determine the bacterial community composition, egg masses from two natural populations in Belgium and Italy were examined. Subsequently, microbial community establishment was assessed by studying stinkbug eggs of different ages obtained from laboratory strains (unlaid eggs collected from the ovaries, eggs less than 24 h old, and eggs collected 4 days after oviposition). Both the external and internal egg-associated microbiomes were analyzed by investigating egg washes and surface-sterilized washed eggs, respectively. Eggs from the ovaries were completely devoid of bacteria, indicating that egg-associated bacteria were deposited on the eggs during or after oviposition. The bacterial diversity of deposited eggs was very low, with on average 6.1 zero-radius operational taxonomic units (zOTUs) in the external microbiome and 1.2 zOTUs in internal samples of egg masses collected from the field. Bacterial community composition and density did not change significantly over time, suggesting limited bacterial growth. A Pantoea-like symbiont previously found in the midgut of N. viridula was found in every sample and generally occurred at high relative and absolute densities, especially in the internal egg samples. Additionally, some eggs harbored a Sodalis symbiont, which has previously been found in the abdomen of several insects, but so far not in N. viridula populations. We conclude that the egg-associated bacterial microbiome of N. viridula is species-poor and dominated by a few symbionts, particularly the species-specific obligate Pantoea-like symbiont.

Contrasting reproductive traits of competing parasitoids facilitate coexistence on a shared host pest in a biological control perspective.

BACKGROUND: Interspecific competition in insect parasitoids is an important ecological phenomenon that has relevant implications for biological pest control. To date, interspecific intrinsic (=larval) competition has been intensively studied, while investigations on extrinsic (=adult) competition have often lagged behind. In this study we examined the role played by parasitoid reproductive traits and host clutch size on the outcome of extrinsic competition between Trissolcus basalis (Wollaston) and Ooencyrtus telenomicida (Vassiliev), two egg parasitoids of the pest Nezara viridula (L). Laboratory experiments were conducted by allowing both parasitoid species to exploit an egg mass made of 10, 20, 30, or 40 hosts through single or simultaneous releases. Furthermore, under field conditions, egg masses consisting of 10 or 40 hosts were exposed in a tomato crop in order to validate laboratory investigation. RESULTS: The results show that the egg mass size is an important predictor of extrinsic competition in our study system as a higher proportion of T. basalis emerged from large egg masses, while O. telenomicida dominated in small egg masses. Analysis of reproductive traits of parasitoid species indicates that T. basalis has superior abilities in host exploitation compared with O. telenomicida. CONCLUSIONS: We found that contrasting reproductive traits of two competing egg parasitoid species facilitate coexistence on a shared stink bug host. This work also highlights the importance to consider extrinsic competitive interactions between parasitoid species in a biological control perspective. © 2022 The Authors. Pest Management Science published by John Wiley & Sons Ltd on behalf of Society of Chemical Industry.

Detection and monitoring of Drosophila suzukii in raspberry and cherry orchards with volatile organic compounds in the USA and Europe.

Spotted wing drosophila (SWD) causes significant economic loss in fruit crops to growers worldwide. There is immediate need for efficacious and selective monitoring tools that can detect infestations early. Previously, volatile organic compounds derived from apple were studied and a quinary chemical component blend (QB) was identified as the key SWD attractant in a blueberry orchard in the United States. This study's aim was to determine whether previously observed QB efficacy, selectivity, and early detection levels could be attained within raspberry and cherry fields in the USA and Europe. Results demonstrated that sticky trap baited QB dispenser provided earlier SWD detection potential than the usually adopted apple cider vinegar (ACV) trap. The number of SWD captured/trap by QB baited trapping systems was significantly lower than that of the ACV trap. However, percent SWD/trap of QB baited traps was same within cherry. Lower non-target capture will save farmer/grower's labor and time allocated to traps installation and drosophila species identification. Within the USA, SWD selectivity of QB baited liquid traps was consistently greater than sticky trap in raspberry field, suggesting that the QB dispenser can be an alternative to the standard ACV lure and that trap design could improve selectivity further.

Trichoderma harzianum Strain T22 Modulates Direct Defense of Tomato Plants in Response to Nezara viridula Feeding Activity.

Plant growth-promoting fungi belonging to genus Trichoderma are known to help plants when dealing with biotic stressors by enhancing plant defenses. While beneficial effects of Trichoderma spp. against plant pathogens have long been documented, fewer studies have investigated their effect on insect pests. Here, we studied the impact of Trichoderma root colonization on the plant defense responses against stink bug feeding attack. For this purpose, a model system consisting of tomato plant, Solanum lycopersicum cv Dwarf San Marzano, Trichoderma harzianum strain T22 and the southern green stink bug, Nezara viridula, was used. We firstly determined stink bug performance in terms of relative growth rate and survival on tomato plants inoculated by T. harzianum T22. Then, we evaluated relative expression of plant defense-related genes on inoculated plants induced by stink bug feeding. We found evidence that T. harzianum T22 affects tomato defense responses against N. viridula nymphs leading to reduction of growth rate. Our results also showed that T. harzianum T22 enhances plant direct defenses by an early increase of transcript levels of jasmonic acid marker genes. Yet this effect was time-dependent and only detected 8 h after herbivore induction. Taken together, our findings provide better understanding on the mechanisms underlying tomato induced resistance against herbivorous stink bugs.

Identification of Brassicadiene, a Diterpene Hydrocarbon Attractive to the Invasive Stink Bug Bagrada hilaris, from Volatiles of Cauliflower Seedlings, Brassica oleracea var. botrytis.

Brassicadiene, a novel tricyclic diterpene hydrocarbon, was identified by a combination of mass spectrometry, microchemical tests, and analysis of NMR spectra. The compound constitutes >90% of the volatile organic compounds produced by cauliflower seedlings, Brassica oleracea var. botrytis. The invasive stink bug Bagrada hilaris is strongly attracted to brassicadiene, providing a mechanism for this herbivore, which specializes on cruciferous plants, to locate its hosts in a nutrient-rich and vulnerable stage.

The Role of (E)-2-octenyl Acetate as a Pheromone of Bagrada hilaris (Burmeister): Laboratory and Field Evaluation.

The pentatomid bug Bagrada hilaris is a key pest of brassicaceous crops in several areas of the world. Previous studies suggest that mate location of this species is mediated by volatile chemicals produced by males, among which the main compound is (E)-2-octenyl acetate. However, the possible attraction of males, females, and nymphs to this compound has not yet been specifically tested. In this study, we tested the response of B. hilaris females, males, and nymphs to (E)-2-octenyl acetate using an electroantennogram (EAG) and olfactometer in the presence or absence of a host plant. Moreover, (E)-2-octenyl acetate as an attractant lure in field trap bioassays was evaluated. EAG recordings showed that this compound evokes antennal responses in B. hilaris females. Olfactometer behavioral responses showed that females and nymphs were attracted to (E)-2-octenyl acetate, while males showed no attraction. In the field trap bioassays, captures were obtained in traps baited with 5 and 10 mg of (E)-2-octenyl acetate, while in traps loaded with 2 mg and control traps, there were no recorded catches. These results suggest the involvement of (E)-2-octenyl acetate in intraspecific interactions of this species.

Egg parasitoid exploitation of plant volatiles induced by single or concurrent attack of a zoophytophagous predator and an invasive phytophagous pest.

Zoophytophagous insect predators can induce physiological responses in plants by activating defence signalling pathways, but whether plants can respond to facultative phytophagy by recruiting natural enemies remains to be investigated. In Y-tube olfactometer bioassays, using a system including a Vicia faba plant, the zoophytophagous predator Podisus maculiventris and the egg parasitoid Telenomus podisi, we first demonstrated that T. podisi females are attracted by broad bean plants damaged by feeding activity of P. maculiventris and on which host egg masses had been laid, while they are not attracted by undamaged plants or plants damaged by feeding activity alone. In a second experiment, we evaluated the impact of the invasive phytophagous pest Halyomorpha halys on this plant volatile-mediated tritrophic communication. Results showed that the invasive herbivorous adults do not induce plants to recruit the native egg parasitoid, but they can disrupt the local infochemical network. In fact, T. podisi females are not attracted by volatiles emitted by plants damaged by H. halys feeding alone or combined with oviposition activity, nor are they attracted by plants concurrently infested by P. maculiventris and H. halys, indicating the specificity in the parasitoid response and the ability of the invasive herbivore in interrupting the semiochemical communication between plants and native egg parasitoids. To the best of our knowledge, this is the first study showing that zoophytophagous predator attacks induce indirect plant defences similarly to those defence strategies adopted by plants as a consequence of single or concurrent infestations of herbivorous insects.

Foraging behavior of two egg parasitoids exploiting chemical cues from the stink bug Piezodorus guildinii (Hemiptera: Pentatomidae).

Several parasitoids attacking the same host may lead to competition. Adult parasitoids' abilities to find, parasitize and defend hosts determine resource's retention potential. In soybean, two egg parasitoid species, Telenomus podisi and Trissolcus urichi (Hymenoptera: Platygastridae), compete on the egg masses of Piezodorus guildinii (Hemiptera: Pentatomidae) one of the major pest of this crop. We evaluated parasitoid's abilities to exploit hosts' footprints; and parasitoid's behavior when competing for the same host. Both arena residence time and retention time were similar for T. podisi and T. urichi on male or female host footprints. In its turn, T. urichi reentered the area contaminated with P. guildinii more times and staid longer in it than T. podisi. Furthermore, when competing for the same egg mass, each parasitoid species won (was in possession of the host by the end of the experiment) half of the replicates, and the number of times each wasp species contacted host in the first place was similar, without affecting replicate outcome (who ultimately won). Both species started agonistic and non-agonistic encounters. This study provides information about the potential interspecific competition between these parasitoids, which contributes to evaluate the compatibility of multiple natural enemies' biological control programs for stink bugs.

Mating Status of an Herbivorous Stink Bug Female Affects the Emission of Oviposition-Induced Plant Volatiles Exploited by an Egg Parasitoid.

Insect parasitoids are under selection pressure to optimize their host location strategy in order to maximize fitness. In parasitoid species that develop on host eggs, one of these strategies consists in the exploitation of oviposition-induced plant volatiles (OIPVs), specific blends of volatile organic compounds released by plants in response to egg deposition by herbivorous insects. Plants can recognize insect oviposition via elicitors that trigger OIPVs, but very few elicitors have been characterized so far. In particular, the source and the nature of the elicitor responsible of egg parasitoid recruitment in the case of plants induced with oviposition by stink bugs are still unknown. In this paper, we conducted behavioral and molecular investigations to localize the source of the elicitor that attracts egg parasitoids and elucidate the role of host mating in elicitation of plant responses. We used as organism study model a tritrophic system consisting of the egg parasitoid Trissolcus basalis, the stink bug host Nezara viridula and the plant Vicia faba. We found that egg parasitoid attraction to plant volatiles is triggered by extracts coming from the dilated portion of the stink bug spermathecal complex. However, attraction only occurs if extracts are obtained from mated females but not from virgin ones. Egg parasitoid attraction was not observed when extracts coming from the accessory glands (mesadene and ectadene) of male hosts were applied, either alone or in combination to plants. SDS-PAGE electrophoresis correlated with olfactometer observations as the protein profile of the dilated portion of the spermathecal complex was affected by the stink bug mating status suggesting post-copulatory physiological changes in this reproductive structure. This study contributed to better understanding the host location process by egg parasitoids and laid the basis for the chemical characterization of the elicitor responsible for OIPV emission.

Volatile unsaturated hydrocarbons emitted by seedlings of Brassica species provide host location cues to Bagrada hilaris.

Bagrada hilaris Burmeister, is a stink bug native to Asia and Africa and invasive in the United States, Mexico, and more recently, South America. This species can cause serious damage to various vegetable crops in the genus Brassica, with seedlings being particularly susceptible to B. hilaris feeding activity. In this study, the role of volatile organic compounds (VOCs) emitted by seedlings of three Brassica species on the host preference of B. hilaris was evaluated. In dual choice arena and olfactometer bioassays, adult painted bugs preferred B. oleracea var. botrytis and B. napus over B. carinata. Volatiles from B. oleracea seedlings were collected and bioassayed with B. hilaris adults and late stage nymphs, using electroantennographic (EAG) and behavioral (olfactometer) techniques. When crude extracts of the VOCs from B. oleracea var. botrytis seedlings and liquid chromatography fractions thereof were bioassayed, B. hilaris adults and nymphs were attracted to the crude extract, and to a non-polar fraction containing hydrocarbons, whereas there were no responses to the more polar fractions. GC-MS analysis indicated that the main constituents of the non-polar fraction was an as yet unidentified diterpene hydrocarbon, with trace amounts of several other diterpene hydrocarbons. The major diterpene occurred in VOCs from both of the preferred host plants B. oleracea and B. napus, but not in VOCs of B. carinata. Our results suggest that this diterpene, alone or in combination with one or more of the minor compounds, is a key mediator in this insect-plant interaction, and could be a good candidate for use in lures for monitoring B. hilaris in the field.

First extensive characterization of the venom gland from an egg parasitoid: structure, transcriptome and functional role.

The venom gland is a ubiquitous organ in Hymenoptera. In insect parasitoids, the venom gland has been shown to have multiple functions including regulation of host immune response, host paralysis, host castration and developmental alteration. However, the role played by the venom gland has been mainly studied in parasitoids developing in larval or pupal hosts while little is known for parasitoids developing in insect eggs. We conducted the first extensive characterization of the venom of the endoparasitoid Ooencyrtus telenomicida (Vassiliev), a species that develops in eggs of the stink bug Nezara viridula (L.). In particular we investigated the structure of the venom apparatus, its functional role and conducted a transcriptomic analysis of the venom gland. We found that injection of O. telenomicida venom induces: 1) a melanized-like process in N. viridula host eggs (host-parasitoid interaction), 2) impairment of the larval development of the competitor Trissolcus basalis (Wollaston) (parasitoid-parasitoid interaction). The O. telenomicida venom gland transcriptome reveals a majority of digestive enzymes (peptidases and glycosylases) and oxidoreductases (laccases) among the most expressed genes. The former enzymes are likely to be involved in degradation of the host resources for the specific benefit of the O. telenomicida offspring. In turn, alteration of host resources caused by these enzymes may negatively affect the larval development of the competitor T. basalis. We hypothesize that the melanization process induced by venom injection could be related to the presence of laccases, which are multicopper oxidases that belong to the phenoloxidases group. This work contributed to a better understanding of the venom in insect parasitoids and allowed to identify candidate genes whose functional role can be investigated in future studies.

An invasive insect herbivore disrupts plant volatile-mediated tritrophic signalling.

Plants respond to insect attack by emission of volatile organic compounds, which recruit natural enemies of the attacking herbivore, constituting an indirect plant defence strategy. In this context, the egg parasitoid Trissolcus basalis is attracted by oviposition-induced plant volatiles emitted by Vicia faba plants as a consequence of feeding and oviposition by the pentatomid host Nezara viridula. However, this local tritrophic web could be affected by the recent invasion by the alien pentatomid bug Halyomorpha halys, an herbivore that shares the same environments as native pentatomid pests. Therefore, we investigated in laboratory conditions the possible impact of H. halys on the plant volatile-mediated signalling in the local tritrophic web V. faba-N. viridula-T. basalis. We found that T. basalis wasps were not attracted by volatiles induced in the plants by feeding and oviposition activities of H. halys, indicating specificity in the wasps' response. However, the parasitoid attraction towards plant volatiles emitted as a consequence of feeding and oviposition by the associated host was disrupted when host, N. viridula, and non-associated host, H. halys, were concurrently present on the same plant, indicating that invasion by the alien herbivore interferes with established semiochemical webs. These outcomes are discussed in a context of multiple herbivory by evaluating the possible influences of alien insects on local parasitoid foraging behaviour.

Infestation of Broad Bean (Vicia faba) by the Green Stink Bug (Nezara viridula) Decreases Shoot Abscisic Acid Contents under Well-Watered and Drought Conditions.

The response of broad bean (Vicia faba) plants to water stress alone and in combination with green stink bug (Nezara viridula) infestation was investigated through measurement of: (1) leaf gas exchange; (2) plant hormone titres of abscisic acid (ABA) and its metabolites, and of salicylic acid (SA); and (3) hydrogen peroxide (H2O2) content. Furthermore, we evaluated the effects of experimentally water-stressed broad-bean plants on N. viridula performance in terms of adult host-plant preference, and nymph growth and survival. Water stress significantly reduced both photosynthesis (A) and stomatal conductance (gs ), while infestation by the green stink bug had no effects on photosynthesis but significantly altered partitioning of ABA between roots and shoots. Leaf ABA was decreased and root ABA increased as a result of herbivore attack, under both well-watered and water-deprived conditions. Water stress significantly impacted on SA content in leaves, but not on H2O2. However, infestation of N. viridula greatly increased both SA and H2O2 contents in leaves and roots, which suggests that endogenous SA and H2O2 have roles in plant responses to herbivore infestation. No significant differences were seen for green stink bug choice between well-watered and water-stressed plants. However, for green stink bug nymphs, plant water stress promoted significantly lower weight increases and significantly higher mortality, which indicates that highly water-stressed host plants are less suitable for N. viridula infestation. In conclusion two important findings emerged: (i) association of water stress with herbivore infestation largely changes plant response in terms of phytohormone contents; but (ii) water stress does not affect the preference of the infesting insects, although their performance was impaired.

Foraging behaviour of an egg parasitoid exploiting plant volatiles induced by pentatomids: the role of adaxial and abaxial leaf surfaces.

Several phases of herbivorous insect attack including feeding and oviposition are known to induce plant defenses. Plants emit volatiles induced by herbivores to recruit insect parasitoids as an indirect defense strategy. So far, volatiles induced by herbivore walking and their putative role in the foraging behavior of egg parasitoids have not been investigated. In this paper we studied the response of the egg parasitoid Trissolcus basalis toward volatiles emitted by Vicia faba plants as consequence of the walking activity of the host Nezara viridula. Olfactometer bioassays were carried out to evaluate wasp responses to plants in which the abaxial or the adaxial surfaces were subjected to walking or/and oviposition. Results showed that host female walking on the abaxial but not on the adaxial surface caused a repellence effect in T. basalis 24 h after plant treatment. The emission of active volatiles also occurred when the leaf was turned upside-down, indicating a specificity of stress localization. This specificity was supported by the results, which showed that oviposition combined with feeding elicit the induction of plant volatiles, attracting the parasitoid, when the attack occurred on the abaxial surface. Analyses of plant volatile blends showed significant differences between the treatments.

Testing the habituation assumption underlying models of parasitoid foraging behavior.

BACKGROUND: Habituation, a form of non-associative learning, has several well-defined characteristics that apply to a wide range of physiological and behavioral responses in many organisms. In classic patch time allocation models, habituation is considered to be a major mechanistic component of parasitoid behavioral strategies. However, parasitoid behavioral responses to host cues have not previously been tested for the known, specific characteristics of habituation. METHODS: In the laboratory, we tested whether the foraging behavior of the egg parasitoid Trissolcus basalis shows specific characteristics of habituation in response to consecutive encounters with patches of host (Nezara viridula) chemical contact cues (footprints), in particular: (i) a training interval-dependent decline in response intensity, and (ii) a training interval-dependent recovery of the response. RESULTS: As would be expected of a habituated response, wasps trained at higher frequencies decreased their behavioral response to host footprints more quickly and to a greater degree than those trained at low frequencies, and subsequently showed a more rapid, although partial, recovery of their behavioral response to host footprints. This putative habituation learning could not be blocked by cold anesthesia, ingestion of an ATPase inhibitor, or ingestion of a protein synthesis inhibitor. DISCUSSION: Our study provides support for the assumption that diminishing responses of parasitoids to chemical indicators of host presence constitutes habituation as opposed to sensory fatigue, and provides a preliminary basis for exploring the underlying mechanisms.

Lures for red palm weevil trapping systems: aggregation pheromone and synthetic kairomone.

BACKGROUND: The optimisation of the lure is essential for the implementation of trapping systems to control insect pests. In this work, the response of the red palm weevil (RPW), Rhynchophorus ferrugineus Olivier, to increasing emission rates of its aggregation pheromone (ferrugineol) and the efficacy of a convenient synthetic kairomone based on fermentation odours (ethyl acetate and ethanol) have been evaluated in different years and locations along the Mediterranean basin. RESULTS: In general, although capture data and emission had noticeable variability among locations, significantly fewer RPW were captured in pyramidal Picusan® traps with the lowest ferrugineol emission rates tested (0.6-3.8 mg day-1 ). Captures increased rapidly with ferrugineol emission up to 4-5 mg day-1 ; then, higher emission rates did not improve or reduce captures, up to the highest emission rate tested of 50.9 mg day-1 . Thus, there is no evidence of an optimum release rate corresponding to a maximum of RPW catches. Traps baited with the synthetic kairomone (1:3 ethyl acetate/ethanol) captured 1.4-2.2 times more total weevils than traps baited only with ferrugineol. Moreover, in most of the locations, the synthetic blend was at least as effective as the local coattractants used (plant material + molasses). CONCLUSIONS: Ferrugineol emission rate can vary in a wide range without significantly affecting RPW response. Coattractants based on fermenting compounds, ethyl acetate and ethanol, are able to improve the attractant level of ferrugineol and could be employed to replace non-standardised natural kairomones in RPW trapping systems after further optimisation of their proportions and doses. © 2016 Society of Chemical Industry.

Interspecific competition/facilitation among insect parasitoids.

Competition for limited resources is a widespread ecological interaction in animals. In the case of insect parasitoids, species can compete for host resources both at the adult stage as well as at the larval stage. Interspecific competition can play a role in sizing and shaping community structures. In addition of being relevant for basic ecological studies, understanding how interspecific competition between parasitoids affects pest suppression is important for biological control. In this opinion paper we review recent advances in the field of interspecific competition among parasitoids in a biological control perspective. We first discuss adult competition, highlighting which factors are likely to play a role in the outcome of competition when adults interact either directly or indirectly. Then we focus on the interactions occurring between competing larvae that develop within the same host taking also into account the fitness consequences of competition for the larva surviving interspecific competition. We also explore the possibility of interspecific facilitation among parasitoids in those situations in which a given species may benefit from interspecific competition.