Subtle morphological changes in the visual and antennal sensory system of bees and wasps across an urbanisation gradient.

Increased temperature and fragmentation of green spaces in urban areas could drive variations in functional traits of insects. Such morphological shifts may occur for sensory systems, which were previously reported to be prone to change with habitat characteristics in non-urban contexts. Here, we measured traits related to the visual and antennal sensory systems in the bees Halictus scabiosae and Osmia cornuta and the wasp Polistes dominula along an urbanisation gradient within Milan (Italy). We hypothesised that fragmentation could filter for better visual properties, and that higher temperature could filter for fewer thermoreceptors and more olfactory hairs. While controlling for body size, results show subtle but appreciable responses to urbanisation in one or more traits in all species, though not always supporting our hypotheses. O. cornuta shows marginally higher ommatidia density and smaller ommatidia diameter (associated with better visual resolution) in more fragmented sites, as well as marginally fewer thermoreceptors in hotter sites, in agreement with our two predictions. On the other hand, H. scabiosae has marginally smaller antennae and P. dominula has smaller eyes at warmer locations, and the wasp also has smaller antennae and 9th flagellomeres in more fragmented areas. Perhaps higher temperatures accelerate development of sensory system at higher speed than the rest of body in these two species. Our results represent the first evidence of urbanisation effects on the visual and antennal sensory systems of bees and wasps and underline how such effects may involve a much broader bouquet of traits then previously observed.

Recognition of nonself is necessary to activate Drosophila's immune response against an insect parasite.

BACKGROUND: Innate immune responses can be activated by pathogen-associated molecular patterns (PAMPs), danger signals released by damaged tissues, or the absence of self-molecules that inhibit immunity. As PAMPs are typically conserved across broad groups of pathogens but absent from the host, it is unclear whether they allow hosts to recognize parasites that are phylogenetically similar to themselves, such as parasitoid wasps infecting insects. RESULTS: Parasitoids must penetrate the cuticle of Drosophila larvae to inject their eggs. In line with previous results, we found that the danger signal of wounding triggers the differentiation of specialized immune cells called lamellocytes. However, using oil droplets to mimic infection by a parasitoid wasp egg, we found that this does not activate the melanization response. This aspect of the immune response also requires exposure to parasite molecules. The unidentified factor enhances the transcriptional response in hemocytes and induces a specific response in the fat body. CONCLUSIONS: We conclude that a combination of danger signals and the recognition of nonself molecules is required to activate Drosophila's immune response against parasitic insects.

Giant honeybees (Apis dorsata) trade off defensiveness against periodic mass flight activity.

The giant honeybee Apis dorsata (Fabricius, 1793) is an evolutionarily ancient species that builds its nests in the open. The nest consists of a single honeycomb covered with the bee curtain which are several layers of worker bees that remain almost motionless with their heads up and abdomens down on the nest surface, except for the mouth area, the hub between inner- and outer-nest activities. A colony may change this semi-quiescence several times a day, depending on its reproductive state and ambient temperature, to enter the state of mass flight activity (MFA), in which nest organisation is restructured and defense ability is likely to be suppressed (predicted by the mass-flight-suspend-defensiveness hypothesis). For this study, three episode of MFA (mfa1-3) of a selected experimental nest were analysed in a case study with sequences of >60 000 images at 50 Hz, each comprise a short pre-MFA session, the MFA and the post-MFA phase of further 10 min. To test colony defensiveness under normative conditions, a dummy wasp was cyclically presented with a standardised motion programme (Pd) with intervening sessions without such a presentation (nPd). Motion activity at five selected surveillance zones (sz1-5) on the nest were analysed. In contrast to mfa1,2, in mfa3 the experimental regime started with the cyclic presentation of the dummy wasp only after the MFA had subsided. As a result, the MFA intensity in mfa3 was significantly lower than in mfa1-2, suggesting that a colony is able to perceive external threats during the MFA. Characteristic ripples appear in the motion profiles, which can be interpreted as a start signal for the transition to MFA. Because they are strongest in the mouth zone and shift to higher frequencies on their way to the nest periphery, it can be concluded that MFA starts earlier in the mouth zone than in the peripheral zones, also suggesting that the mouth zone is a control centre for the scheduling of MFA. In Pd phases of pre- and postMFA, the histogram-based motion spectra are biphasic, suggesting two cohorts in the process, one remaining at quiescence and the other involved in shimmering. Under MFA, nPd and Pd spectra were typically Gaussian, suggesting that the nest mates with a uniform workload shifted to higher motion activity. At the end of the MFA, the spectra shift back to the lower motion activities and the Pd spectra form a biphasic again. This happens a few minutes earlier in the peripheral zones than in the mouth zone. Using time profiles of the skewness of the Pd motion spectra, the mass-flight-suspend-defensiveness hypothesis is confirmed, whereby the inhibition of defense ability was found to increase progressively during the MFA. These sawtooth-like time profiles of skewness during MFA show that defense capability is recovered again quite quickly at the end of MFA. Finally, with the help of the Pd motion spectra, clear indications can be obtained that the giant honeybees engage in a decision in the sense of a tradeoff between MFA and collective defensiveness, especially in the regions in the periphery to the mouth zone.

Immune response in paper wasp workers: Task matters more than age.

Workers of social hymenopterans (ants, bees and wasps) display specific tasks depending on whether they are younger or older. The relative importance of behavior and age in modulating immune function has seldom been addressed. We compared the strength of encapsulation-melanization immune response (hereafter melanotic encapsulation) in paper wasps displaying age polyethism or experimentally prevented from behavioral specialization. Foragers of Polybia paulista had higher melanotic encapsulation than guards, regardless of their age. Nevertheless, melanotic encapsulation decreased with age when wasps were prevented from behavioral specialization. Thus, in this species, worker melanotic encapsulation seems more sensitive to task than age. Foraging is considered one of the riskier behaviors in terms of pathogen exposure, so upregulating melanotic encapsulation in foragers can possibly improve both individual and colony-level resistance against infections.

Effect of egg production dynamics on the functional response of two parasitoids.

Functional response describes the number of hosts attacked by a parasitoid in relation to host densities and plays an important role by connecting behavioral-level processes with community-level processes. Most functional response studies were carried out using simple experimental designs where the insects were confined to a plain and small arena with different host densities during a fixed period of time. With these designs, other factors that might affect the functional response of parasitoids were not analyzed, such as fecundity, age, and experience. We proposed a series of latent-variables Markovian models that comprised an integrated approach of functional response and egg production models to estimate the realized lifetime reproductive success of parasitoids. As a case study, we used the parasitoids Anagyrus cachamai and A. lapachosus (Hymenoptera: Encyrtidae), two candidate agents for neoclassical biocontrol of the Puerto Rican cactus pest mealybug, Hypogeococcus sp. (Hemiptera: Pseudococcidae). The tested species were assessed according to their physiology and prior experience. We estimated the number of mature eggs after emergence, egg production on the first day, egg production rate, the proportion of eggs resorbed, egg resorption threshold, and egg storage capacity. Anagyrus cachamai and A. lapachosus both presented a type III functional response. However, the two parasitoids behaved differently; for A. cachamai, the number of parasitized hosts decreased with female age and depended on the number of mature eggs that were available for oviposition, whereas A. lapachosus host parasitism increased with female age and was modulated by its daily egg load and previous experience. The methodology presented may have large applicability in pest control, invasive species management, and conservation biology, as it has the potential to increase our understanding of the reproductive biology of a wide variety of species, ultimately leading to improved management strategies.

A test of trade-offs in dispersal and reproduction within and between a sister species pair of specialist insect herbivores.

Understanding the drivers of trade-offs among traits is vital for comprehending the evolution and maintenance of trait variation. Theoretical frameworks propose that evolutionary mechanisms governing trade-offs frequently exhibit a scale-dependent nature. However, empirical tests of whether trade-offs exhibited across various biological scales (i.e. individuals, populations, species, genera, etc.) remains scarce. In this study, we explore trade-off between dispersal and reproductive effort among sympatric sister species of wasps in the genus Belonocnema (Hymenoptera: Cynipini: Cynipidae) that form galls on live oaks: B. fossoria, which specializes on Quercus geminata, and B. treatae, which specializes on Q. virginiana. Specifically, our results suggest that B. fossoria has evolved reduced flight capability and smaller wings, but a larger abdomen and greater total reproductive effort than B. treatae, which has larger wings and is a stronger flier, but has a smaller abdomen and reduced total reproductive effort. These traits and the relationships among them remain unchanged when B. fossoria and B. treatae are transplanted and reared onto the alternative host plant, suggesting that trait divergence is genetically based as opposed to being a plastic response to the different rearing environments. However, when looking within species, we found no evidence of intraspecific trade-offs between wing length and reproductive traits within either B. fossoria or B. treatae. Overall, our results indicate that observed trade-offs in life history traits between the two gall former species are likely a result of independent adaptations in response to different environments as opposed to the amplified expression of within species intrinsic tradeoffs.

Oviposition behavior of the quasi-gregarious parasitoid, Ooencyrtus kuvanae (Hymenoptera: Encyrtidae).

The parasitoid wasp, Ooencyrtus kuvanae (Howard) (Hymenoptera: Encyrtidae), is a natural enemy of the spongy moth, a significant forest pest in North America. We investigated the oviposition behavior of O. kuvanae females on spongy moth egg masses by (i) presenting female parasitoids with a single spongy moth egg mass that was replaced every day, 2nd day, 4th day, 8th day, or 16th day (which is the total length of the oviposition period) and (ii) presenting female parasitoids with 1, 2, 4, or 8 egg masses at a time. Offspring developmental length ranged from 18 to 24 days. On average, male offspring exhibited faster developmental times, emerging approximately 1 day ahead of females. The amount of time that adult females spent on an egg mass affected the number of parasitized eggs. Specifically, more offspring emerged in the 4-, 8-, and 16-day treatments than in scenarios involving daily or every second-day egg mass replacement. The percentage of male offspring decreased as the number of egg masses presented to females increased. Interestingly, the total number of female offspring remained constant, but the number of male offspring decreased with an increase in the number of egg masses and time spent by the parent within a patch. The observed sexual dimorphism in development time, the influence of resource availability on offspring sex ratios, and flexible oviposition patterns illustrate the adaptability of O. kuvanae in response to varying conditions. These insights have implications for our understanding of parasitoid-host interactions and their potential role in biological control strategies.

Ozone alters the chemical signal required for plant - insect pollination: The case of the Mediterranean fig tree and its specific pollinator.

Tropospheric ozone (O3) is likely to affect the chemical signal emitted by flowers to attract their pollinators through its effects on the emission of volatile organic compounds (VOCs) and its high reactivity with these compounds in the atmosphere. We investigated these possible effects using a plant-pollinator interaction where the VOCs responsible for pollinator attraction are known and which is commonly exposed to high O3 concentration episodes: the Mediterranean fig tree (Ficus carica) and its unique pollinator, the fig wasp (Blastophaga psenes). In controlled conditions, we exposed fig trees bearing receptive figs to a high-O3 episode (5 h) of 200 ppb and analyzed VOC emission. In addition, we investigated the chemical reactions occurring in the atmosphere between O3 and pollinator-attractive VOCs using real-time monitoring. Finally, we tested the response of fig wasps to the chemical signal when exposed to increasing O3 mixing ratios (0, 40, 80, 120 and 200 ppb). The exposure of the fig tree to high O3 levels induced a significant decrease in leaf stomatal conductance, a limited change in the emission by receptive figs of VOCs not involved in pollinator attraction, but a major change in the relative abundances of the compounds among pollinator-attractive VOCs in O3-enriched atmosphere. Fig VOCs reacted with O3 in the atmosphere even at the lowest level tested (40 ppb) and the resulting changes in VOC composition significantly disrupted the attraction of the specific pollinator. These results strongly suggest that current O3 episodes are probably already affecting the interaction between the fig tree and its specific pollinator.

Flourishing with sugars - following the fate of parasitoids in the field.

When flowers, plants bearing extrafloral nectaries, or sugar sprays are added to agroecosystems, parasitoids are expected to feed, thereby improving biological control. This paper reviews studies where sugar-feeding of field-collected parasitoids were monitored via biochemical assays. When examined, trends in parasitoid abundance, energetic reserves, longevity, and per capita fecundity are also followed. Starved parasitoids readily feed on sugar sources in the field, and more individuals collected near sugar sources are categorized as 'fed' when sugar is otherwise limited in the agroecosystem. When sugar is not limited (i.e. honeydew prevalent), trends are not as clear. Some studies show improved fecundity and parasitism, while other studies show inconsistent trends between parasitoid feeding, abundance, longevity, and parasitism, with some responses improved but not others. Future research could address the dispersal or resting behavior of wasps following feeding since it can influence eventual biological control, and consider whether field sampling methods might over-/underestimate feeding.

Parasitoid cues modulate Drosophila germline development and stem cell proliferation.

Environmental factors influence an organism's reproductive ability by regulating germline development and physiology. While the reproductive adaptations in response to extrinsic stress cues offer fitness and survival advantages to individuals, the mechanistic understanding of these modifications remains unclear. Here, we find that parasitoid wasps' stress signaling regulates Drosophila melanogaster oogenesis. We show that fruit flies dwelling in the wasp-infested area elevate their fecundity, and the observed reproductive response is specific to Pachycrepoideus sp., a pupal parasitoid wasp. Pachycrepoideus-specific olfactory and visual cues recruit the signaling pathways that promote germline stem cell proliferation and accelerate follicle development, increasing egg production in Drosophila females. Downregulation of signaling engaged in oocyte development by shifting flies to a non-wasp-infested environment increases apoptosis of the developing follicles. Thus, this study establishes host germline responsiveness to parasitoid-specific signals and supports a predator strategy to increase hosts for infection.

Tritrophic effects mediate host suitability for two parasitoids of the carob moth (Lepidoptera: Pyralidae), infesting pistachio kernels.

The carob moth, Ectomyelois ceratoniae (Zeller) (Lepidoptera: Pyralidae), is an important pest of pistachio causing direct damage to nuts during ripening, and in storage. We examined the tritrophic effects of 5 pistachio cultivars, representing the full range of kernel suitability for E. ceratoniae, on 2 larval parasitoids, Habrobracon hebetor (Say) (Hymenoptera: Braconidae), a gregarious, synovigenic, idiobiont, ectoparasitoid, and Venturia canescens Gravenhorst (Hymenoptera: Ichneumonidae), a solitary, pro-ovigenic, koinobiont, endoparasitoid. The suitability of pistachio cultivars for development and reproduction of the 2 parasitoids was closely correlated, despite their divergent lifestyles and developmental patterns, and paralleled variation in kernel suitability for host larvae. Although kernels of the most resistant cultivar reduced demographic parameters of the parasitoids, the potential impact on numerical responses was minor. Successful biological control of carob moth within the vulnerable period of nut ripening will likely depend on the functional responses of earlier generations of adult parasitoids emerging from other fruit and nut crops, or those released in an augmentation program.

Comparative transcriptome and proteome reveal the unique genes and proteins of female parasitic wasps, Lysiphlebia japonica Ashmead.

BACKGROUND: Lysiphlebia japonica Ashmead (Hymenoptera, Braconidae) is an endophagous parasitoid wasp and its host, Aphis gossypii Glover (Hemiptera, Aphididae) is a major cotton pest. L. japonica affects the growth and fatty acid metabolism of cotton aphids after parasitization and has been widely used as a biocontrol agent. However, there are currently few reports about the molecular characteristics of L. japonica, especially the differences between male and female. RESULTS: In this study, using transcriptome and proteome analysis of the abdomen of female and male parasitic wasps, respectively, we obtained a total of 27,169 DEGs and 1,194 DEPs, then a total of 909 positively correlated high-expression proteins and genes were obtained by combined omics analysis. Subsequently, 20 differentially expressed abdomen specific proteins were selected for validation by RT-qPCR and Multiple Reaction Monitoring (MRM) protein verification. The result of RT-qPCR demonstrated that all 20 genes were highly expressed in the abdomen of females, and five target proteins with unique peptide fragments and identification profiles were identified by MRM, which were venom protease, tropomyosin, lipase member I, venom serine carboxypeptidase and calreticulin, respectively. CONCLUSION: Overall, these results provided molecular resources for the differences between males and females in L. japonica and the screened 20 abdomen specific proteins were verified to demonstrate the validity of the data, which offered important molecular data resources for further studies on the related functional genes of parasitic wasps and the mechanism of parasitoids regulating the growth of aphids. © 2023 Society of Chemical Industry.

Effects of root inoculation of entomopathogenic fungi on olfactory-mediated behavior and life-history traits of the parasitoid Aphidius ervi (Haliday) (Hymenoptera: Braconidae).

BACKGROUND: Although most biological control programs use multiple biological agents to manage pest species, to date only a few programs have combined the use of agents from different guilds. Using sweet pepper (Capsicum annuum L.), the entomopathogenic fungus Akanthomyces muscarius ARSEF 5128, the tobacco peach aphid Myzus persicae var. nicotianae and the aphid parasitoid Aphidius ervi as the experimental model, we explored whether root inoculation with an entomopathogenic fungus is compatible with parasitoid wasps for enhanced biocontrol of aphids. RESULTS: In dual-choice behavior experiments, A. ervi was significantly attracted to the odor of M. persicae-infested C. annuum plants that had been inoculated with A. muscarius, compared to noninoculated infested plants. There was no significant difference in attraction to the odor of uninfested plants. Myzus persicae-infested plants inoculated with A. muscarius emitted significantly higher amounts of indole, (E)-nerolidol, (3E,7E)-4,8,12-trimethyltrideca-1,3,7,11-tetraene and one unidentified terpene compared to noninoculated infested plants. Coupled gas chromatography-electroantennography, using the antennae of A. ervi, confirmed the physiological activity of these elevated compounds. Inoculation of plants with A. muscarius did not affect parasitism rate nor parasitoid longevity, but significantly increased the speed of mummy formation in parasitized aphids on fungus-inoculated plants. CONCLUSION: Our data suggest that root inoculation of C. annuum with A. muscarius ARSEF 5128 alters the olfactory-mediated behavior of parasitoids, but has little effect on parasitism efficiency or life-history parameters. However, increased attraction of parasitoids towards M. persicae-infested plants when inoculated by entomopathogenic fungi can accelerate host localization and hence improve biocontrol efficacy. © 2023 Society of Chemical Industry.

Identification of floral volatiles from Fagopyrum esculentum that attract Cotesia vestalis with potentially better biocontrol efficacy against Plutella xylostella.

BACKGROUND: Nectar plants provide extra nourishment for parasitoids, which can utilize floral volatiles to locate nectar-rich flowers. A promising strategy is to screen potential floral species based on the wasps' olfactory preferences for nectar sources, and to ensure their suitability for both natural enemies and targeted pests. Cotesia vestalis (Haliday) is a dominant parasitoid of the oligophagous pest Plutella xylostella, which poses a significant threat to cruciferous vegetables globally. However, the chemical cues in plant-parasitoid complexes mediating Cotesia vestalis to locate nectar food resources and the positive effect of nectar plants on the Cotesia vestalis population are poorly understood. RESULTS: The results showed that Fagopyrum esculentum was the most attractive plant that attracted Cotesia vestalis, not Plutella xylostella in 44 flowering plants from 19 families. 1,2-Diethyl benzene and 1,4-diethyl benzene, identified from the floral volatiles from F. esculentum in full bloom, were found to elicit dose-dependent electrophysiological responses and attract Cotesia vestalis adults, demonstrating their potential as semiochemicals. Moreover, the age-stage, two-sex life table revealed that feeding on nectar food increased the efficacy of Cotesia vestalis adults against Plutella xylostella. CONCLUSION: In summary, the findings provide insights into the chemical ecology of plant-parasitoid complexes and support the potential use of F. esculentum as insectary plants in habitat manipulation against Plutella xylostella by supplying natural nectar food for the Cotesia vestalis population. Our results suggest an attract and reward strategy based on an attractant for Cotesia vestalis to control Plutella xylostella, or the development of volatile-based artificial food for Cotesia vestalis. © 2023 Society of Chemical Industry.

Parasitoid-induced changes in metabolic rate and feeding activity of the emerald ash borer (Coleoptera: Buprestidae): implications for biological control.

Parasitoid-host interactions form the foundation of biological control strategies against many agriculture and forest insect pests. The emerald ash borer (EAB), Agrilus planipennis (Coleoptera: Buprestidae), is a serious invasive pest of ash (Fraxinus spp.) trees in North America. Tetrastichus planipennisi (Hymenoptera: Eulophidae) is a gregarious, koinobiont endoparasitoid, attacking late (3rd to 4th) instars of EAB larvae, which feed in the live phloem of ash trunks or branches, making serpentine-like galleries filled with larval frass. In the present study, we tested the hypothesis that T. planipennisi regulates the host metabolism and feeding activity to optimize its offspring development and fitness. We first compared the respiration rate of parasitized and unparasitized host larvae at different times after parasitism, and then measured feeding activity of both parasitized and unparasitized host larvae inside their feeding galleries. Although parasitized host larvae increased metabolic rate and feeding activity in the first few days of parasitism, T. planipennisi parasitism induced an overall reduction of the metabolic rate and decrease in feeding activity of parasitized host larvae over their development period. In addition, there was a negative relationship between feeding activity of parasitized hosts and brood sizes of the parasitoid progeny-i.e., the more parasitoid progeny a host larva received, the less feeding activity the host had. These findings suggest that T. planipennisi has limited ability to optimize its offspring development and fitness through regulations of the host metabolism and feeding activity and its parasitism reduces feeding damage of parasitized EAB larvae to infested ash trees.

Sugar responsiveness could determine foraging patterns in yellowjackets.

Sympatric-related species often exhibit resource partitioning. This can occur through different mechanisms, such as behavioral, morphological, and sensory variations, leading to qualitative, temporal, or spatial differences in resource exploitation, such as consuming different types of food. Sensory-based niche partitioning could be the underlying mechanism through which closely related species effectively reduce niche overlap. Here we ask whether variations in sensory responses to carbohydrates could reflect differences in the foraging patterns of two Vespula species present in Patagonia. For this, we established (i) the response thresholds toward carbohydrate solutions of foraging V. germanica and V. vulgaris in the laboratory, (ii) the sugar concentration of foraged carbohydrates in the field, and (iii) possible effects of incoming sugar concentration and performance at individual and colony levels. Results indicate a higher sucrose response threshold in V. germanica than V. vulgaris. Field results indicate that higher carbohydrate concentrations foraged by V. germanica, with 57% of V. germanica foragers returning with concentrations above 50% w/w, while only 23% of V. vulgaris foragers did so. These differences in sucrose sensitivity and foraging patterns positively correlate with colony size, irrespective of the species. Our results suggest that competition could be reduced in these closely related invasive social wasp species through sensory differences in their sugar perception levels, which would lead to them foraging different carbohydrate sources. This study suggests that sensory niche partitioning could promote species coexistence in these social wasps.

Extracting Venom from the Parasitoid Wasp Trichogramma dendrolimi using an Artificial Host.

Parasitoid wasps are a diverse group of hymenopteran insects that serve as invaluable resources for pest biocontrol. To ensure successful parasitism, parasitoid wasps inject venom into their hosts to suppress their hosts' immunity, modulate hosts' development, metabolism, and even behavior. With over 600,000 estimated species, the diversity of parasitoid wasps surpasses that of other venomous animals, such as snakes, cone snails, and spiders. Parasitoid wasp venom is an underexplored source of bioactive molecules with potential applications in pest control and medicine. However, collecting parasitoid venom is challenging due to the inability to use direct or electrical stimulation and the difficulty in dissection because of their small size. Trichogramma is a genus of tiny (~0.5 mm) egg parasitoid wasps that are widely used for the biological control of lepidopteran pests in both agriculture and forests. Here, we report a method for extracting venom from T. dendrolimi using artificial hosts. These artificial hosts are created with polyethylene film and amino acid solutions and then inoculated with Trichogramma wasps for parasitism. The venom was subsequently collected and concentrated. This method enables the extraction of large amounts of Trichogramma venom while avoiding contamination from other tissues caused by dissection, a common issue in venom reservoir dissection protocols. This innovative approach facilitates the study of Trichogramma venom, paving the way for new research and potential applications.

Indirect plant-mediated interactions between heterospecific parasitoids that develop in different caterpillar species.

Parasitoids induce physiological changes in their herbivorous hosts that affect how plants respond to herbivory. The signature of parasitoids on induced plant responses to feeding by parasitized herbivores indirectly impacts insect communities interacting with the plant. The effect may extend to parasitoids and cause indirect interaction between parasitoids that develop inside different herbivore hosts sharing the food plant. However, this type of interactions among parasitoid larvae has received very little attention. In this study, we investigated sequential and simultaneous plant-mediated interactions among two host-parasitoid systems feeding on Brassica oleracea plants: Mamestra brassicae parasitized by Microplitis mediator and Pieris rapae parasitized by Cotesia rubecula. We measured the mortality, development time, and weight of unparasitized herbivores and performance of parasitoids that had developed inside the two herbivore species when sharing the food plant either simultaneously or sequentially. Plant induction by parasitized or unparasitized hosts had no significant effect on the performance of the two herbivore host species. In contrast, the two parasitoid species had asymmetrical indirect plant-mediated effects on each other's performance. Cotesia rubecula weight was 15% higher on plants induced by M. mediator-parasitized hosts, compared to control plants. In addition, M. mediator development time was reduced by 30% on plants induced by conspecific but not heterospecific parasitoids, compared to plants induced by its unparasitized host. Contrary to sequential feeding, parasitoids had no effect on each other's performance when feeding simultaneously. These results reveal that indirect plant-mediated interactions among parasitoid larvae could involve any parasitoid species whose hosts share a food plant.

Chemical Resemblance of Egg Surface Compounds and Dufour's Gland in Two Neotropical Polistinae Wasps Polistes versicolor (Olivier) and Mischocyttarus metathoracicus (de Saussure, 1854).

Chemical communication plays a major role in regulating social dynamics in social insect colonies. The most studied class of chemical compounds are the cuticular hydrocarbons (CHCs), compounds with high molecular weight that cover the insect body. CHCs are used in nestmate recognition and to signal reproductive status. Brood, in the form of larvae and eggs, is known to participate in chemical communication and social dynamics by performing hunger behaviour and inducing interaction with adults and conferring nest and maternity identity. CHCs of adults and egg surface compounds are similar in composition in social insect species. The main source of egg compounds is proposed to be Dufour's gland, an accessory reproductive gland found in several Hymenoptera females. There is still a lack of information about the level of similarity among CHCs, compounds of egg surface and Dufour's gland for several wasp species, which could provide correlational evidence about the origins of egg-marking compounds. Thus, we investigated whether egg surface compounds were more similar to CHCs or Dufour's gland secretions in two Neotropical primitively eusocial wasp species, Polistes versicolor (Olivier) and Mischocyttarus metathoracicus (de Saussure, 1854). As expected, there was a higher chemical similarity between eggs and Dufour's gland secretions in both studied species, supporting the hypothesis that this gland is the source of chemical compounds found over the eggs in these two primitively eusocial species.

Evolution of the neuronal substrate for kin recognition in social Hymenoptera.

In evolutionary terms, life is about reproduction. Yet, in some species, individuals forgo their own reproduction to support the reproductive efforts of others. Social insect colonies for example, can contain up to a million workers that actively cooperate in tasks such as foraging, brood care and nest defence, but do not produce offspring. In such societies the division of labour is pronounced, and reproduction is restricted to just one or a few individuals, most notably the queen(s). This extreme eusocial organisation exists in only a few mammals, crustaceans and insects, but strikingly, it evolved independently up to nine times in the order Hymenoptera (including ants, bees and wasps). Transitions from a solitary lifestyle to an organised society can occur through natural selection when helpers obtain a fitness benefit from cooperating with kin, owing to the indirect transmission of genes through siblings. However, this process, called kin selection, is vulnerable to parasitism and opportunistic behaviours from unrelated individuals. An ability to distinguish kin from non-kin, and to respond accordingly, could therefore critically facilitate the evolution of eusociality and the maintenance of non-reproductive workers. The question of how the hymenopteran brain has adapted to support this function is therefore a fundamental issue in evolutionary neuroethology. Early neuroanatomical investigations proposed that social Hymenoptera have expanded integrative brain areas due to selection for increased cognitive capabilities in the context of processing social information. Later studies challenged this assumption and instead pointed to an intimate link between higher social organisation and the existence of developed sensory structures involved in recognition and communication. In particular, chemical signalling of social identity, known to be mediated through cuticular hydrocarbons (CHCs), may have evolved hand in hand with a specialised chemosensory system in Hymenoptera. Here, we compile the current knowledge on this recognition system, from emitted identity signals, to the molecular and neuronal basis of chemical detection, with particular emphasis on its evolutionary history. Finally, we ask whether the evolution of social behaviour in Hymenoptera could have driven the expansion of their complex olfactory system, or whether the early origin and conservation of an olfactory subsystem dedicated to social recognition could explain the abundance of eusocial species in this insect order. Answering this question will require further comparative studies to provide a comprehensive view on lineage-specific adaptations in the olfactory pathway of Hymenoptera.