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Understanding the major mechanisms that allow an alien species to become invasive is crucial for limiting the impact of such species, and individual immunity seems to be a crucial trait for their large range colonization when introduced to a new area. Indeed, a high resistance to pathogens and parasites could favour the establishment and proliferation of an alien species in a newly colonized region. A striking example of recent successful invasion is represented by the Asian hornet, Vespa velutina nigrithorax, which has invaded Western Europe threatening bees and beekeeping activities. Investigating the biology of this species is fundamental to understand how V. velutina nigrithorax has become invasive in Europe and this knowledge could be crucial for limiting its impact. Here, by performing bacterial challenge bioassays, we compare the immune competence of different castes and sex of a population of the invasive Asian hornet, V. velutina nigrithorax, and of the native European hornet, V. crabro. Our results demonstrate that the two species differ in their immunocompetence with respect to caste: a higher level of immunocompetence was found in the reproductive females of the Asian hornet which were resistant to bacterial challenge when compared to workers and males of both species and to reproductive females of the native European hornet. The high pathogen resistance in reproductive females of the Asian hornet, might therefore represent a key factor contributing to the ecological success and spread of this invader. [ABSTRACT FROM AUTHOR]

Parasites and pathogens can follow different patterns of infection depending on the host developmental stage or sex. In fact, immune function is energetically costly for hosts and trade‐offs exist between immune defenses and life history traits as growth, development and reproduction and organisms should thus optimize immune defense through their life cycle according to their developmental stage. Identifying the most susceptible target and the most virulent pathogen is particularly important in the case of insect pests, in order to develop effective control strategies targeting the most vulnerable individuals with the most effective control agent. Here, we carried out laboratory tests to identify the most susceptible target of infection by infecting different stages of the red palm weevil Rhynchophorus ferrugineus (larvae, pupae, male, and female adults) with both a generic pathogen, antibiotic‐resistant Gram‐negative bacteria Escherichia coli XL1‐Blue, and two specific strains of entomopathogenic nematodes (EPNs), Steinernema carpocapsae ItS‐CAO1 and Heterorhabditis bacteriophora ItH‐LU1. By evaluating bacterial clearance, host mortality and parasite progeny release, we demonstrate that larvae are more resistant than adults to bacterial challenge and they release less EPNs progeny after infection despite a higher mortality compared to adults. Considering the two EPN strains, S. carpocapsae was more virulent than H. bacteriophora both in terms of host mortality and more abundant progeny released by hosts after death. The outcomes attained with unspecific and specific pathogens provide useful information for a more efficient and sustainable management of this invasive pest. [ABSTRACT FROM AUTHOR]

Social recognition represents the foundation of social living. To what extent social recognition is hard-wired by early-life experience or flexible and influenced by social context of later life stages is a crucial question in animal behaviour studies. Social insects have represented classic models to investigate the subject, and the acknowledged idea is that relevant information to create the referent template for nest-mate recognition (NMR) is usually acquired during an early sensitive period in adult life. Experimental evidence, however, highlighted that other processes may also be at work in creating the template and that such a template may be updated during adult life according to social requirements. However, currently, we lack an ad hoc experiment testing the alternative hypotheses at the basis of NMR ontogeny in social insects. Thus, to investigate the mechanisms underlying the ontogeny of NMR in Polistes wasps, a model genus in recognition studies, and their different role in determining recognition abilities, we subjected Polistes dominula workers to different olfactory experiences in different phases of their life before inserting theminto the social environment of a novel colony and testing them in recognition bioassays. Our results show that workers develop their NMR abilities based on their social context rather than through pre-imaginal and early learning or self-referencing. Our study demonstrates that the social context represents the major component shaping recognition abilities in a social wasp, therefore shedding new light on the ontogeny of recognition in paper wasps and prompting the reader to rethink about the traditional knowledge at the basis of the recognition in social insects. [ABSTRACT FROM AUTHOR]

Eusocial insect colonies represent some of the most extreme examples of specialized division of labor. Ageing in workers is often associated with a temporal polyethism in the tasks performed both inside and outside the colony. Such behavioral transition is sometimes linked to a gradual reduction in individual immunity. Here, we studied the immune ability of Apis mellifera guard bees, which represent an intermediate stage between house bees working inside the nest and foragers collecting resources outside, to assess if their specific task is associated with an immune specialization. Through immune challenge with Gram-negative bacteria Escherichia coli, we compared the guards ability to clear bacterial cells from their haemolymph with respect to house bees and foragers. Our findings demonstrate that guards do not show an immune specialization linked to their task but seem to represent a transition also in terms of immunity, since their anti-bacterial response appears intermediate between house bees and foragers. [ABSTRACT FROM AUTHOR]

In the early stage of the invasion process, alien species may face costs linked to pioneer effect due to genetic bottleneck, drift and the consequential inbreeding depression. Thus, introduced species that show an immediate exponential growth in their invasive population should have some mechanisms to reduce such costs minimising the chance of inbreeding or allowing them to cope with them. The yellow-legged hornet Vespa velutina nigrithorax has been spotted in France in 2004; since then, the species has been invading Europe with a relentless pace. In their native range, males and reproductive females of a Chinese non-invasive sub-species of V. velutina seem to leave their nests to search for unrelated partners. However, previous studies showed a low genetic diversity and a high rate of diploid males in colonies of the invasive population, suggesting that mating could occur inside nests, where males should be able to discriminate between reproductive gynes and sterile workers. Here, we used laboratory behavioural assays to investigate the mating preferences of yellow-legged hornet males from the recent invasive population in Italy. We assessed the importance of nestmateship and female morpho-physiological traits, likely indicators of caste, in determining male attraction towards potential partners. Our results demonstrate that males are more attracted to bigger females with more abundant fat storage, good indicators of female reproductive caste in wasps, regardless of nestmateship. Our findings represent a first step in understanding the reproductive biology of V. velutina nigrithorax in its invasive range, providing a framework for future research in the field to prevent or reduce the species expansion. [ABSTRACT FROM AUTHOR]

The parasitic insect Xenos vesparum induces noticeable behavioral and physiological changes—e.g. castration—in its female host, the paper wasp Polistes dominula: parasitized putative workers avoid any colony task and desert the colony to survive in the nearby vegetation, like future queens and males do. In this long-term observational study, we describe the spectacular attraction of parasitized workers towards trumpet creeper bushes (Campsis radicans) in early-summer. Two thirds of all wasps that we sampled on these bushes were parasitized, whereas the parasite prevalence was much lower in our study area and most wasps sampled on other nearby flowering bushes were non-parasitized. First, we describe the occurrence and consistency of this phenomenon across different sites and years. Second, we evaluate the spatial behavior of parasitized wasps on C. radicans bushes, which includes site-fidelity, exploitation and defense of rich extra-floral nectaries on buds and calices. Third, we record two critical steps of the lifecycle of X. vesparum on C. radicans: the parasite’s mating and a summer release of parasitic larvae, that can infect larval stages of the host if transported to the host’s nest. In a nutshell, C. radicans bushes provide many benefits both to the parasite X. vesparum and to its host: they facilitate the parasite’s mating and bivoltine lifecycle, a phenomenon never described before for this parasite, while, at the same time, they provide the wasp host with shelter inside trumpet flowers and extrafloral gland secretions, thus likely enhancing host survival and making it a suitable vector for the infection. [ABSTRACT FROM AUTHOR]

Differences in male and female phenotypes are often the results of sexual selection. Over the years, a vast number of studies investigated how and why the 2 sexes differ in their physical appearance, reaching the conclusion that peculiar visual traits, signals, or ornaments usually evolve under the pressure of sexual selection to mediate intrasexual competition and mate choice. In social contexts, however, males and females can hold different roles and social interactions may depend on the individual gender. The female-dominated hymenopteran societies represent a fascinating scenario to investigate recognition and communication among individuals of different genders outside a common sexual selection framework because sterile female workers typically do not mate and are not attracted to males. Here, we used laboratory bioassays (lure presentation experiments) to evaluate the ability of Polistes dominula workers to discriminate between individuals of the 2 genders, investigating the relevance of the chemical and visual cues potentially involved in such process. Our results showed that P. dominula workers are able to discriminate between the sexes and visual cues rather than chemical ones are responsible for such discrimination. [ABSTRACT FROM AUTHOR]

Males are under different selective pressures than females, which results in differences in the physiology of the two sexes to maximize their fitness. In terms of immunity, males are typically considered as the 'sicker sex', where immunocompetence is reduced to favour increased reproductive output. However, male social Hymenoptera are also haploid and therefore lack allelic variation at the individual level, which can also lead to reduced immunocompetence. Over the last decade, several studies have provided evidence for a higher susceptibility to disease in males of social Hymenoptera, without clarifying whether this susceptibility was a direct consequence of their haploid condition or the result of a 'live hard, die young' overall evolutionary strategy. In the present study, we used an experimental approach of bacterial challenge to test the immune response of males and females in two species of social Hymenoptera (honey bees, Apis mellifera; paper wasps, Polistes dominula), where males show very different life-history traits. Drones benefit from colony protection for most of their life, whereas P. dominula males leave their colonies and have to survive for weeks at leks. If the haploid condition is responsible for a higher susceptibility in males, we should expect a lower immune response in males of both species compared to females. Conversely, if the immunocompetence depends on the life-history traits of males, an opposite trend is expected in males of the two species. Our results do not support the 'haploid susceptibility hypothesis' but are in accordance with the different life history of males from the two species. © 2014 The Linnean Society of London, Biological Journal of the Linnean Society, 2015, 114, 274-278. [ABSTRACT FROM AUTHOR]

The importance of early experience in animals’ life is unquestionable, and imprinting-like phenomena may shape important aspects of behaviour. Early learning typically occurs during a sensitive period, which restricts crucial processes of information storage to a specific developmental phase. The characteristics of the sensitive period have been largely investigated in vertebrates, because of their complexity and plasticity, both in behaviour and neurophysiology, but early learning occurs also in invertebrates. In social insects, early learning appears to influence important social behaviours such as nestmate recognition. Yet, the mechanisms underlying recognition systems are not fully understood. It is currently believed that Polistes social wasps are able to discriminate nestmates from non-nestmates following the perception of olfactory cues present on the paper of their nest, which are learned during a strict sensitive period, immediately after emergence. Here, through differential odour experience experiments, we show that workers of Polistes dominula develop correct nestmate recognition abilities soon after emergence even in absence of what have been so far considered the necessary cues (the chemicals spread on nest paper). P. dominula workers were exposed for the first four days of adult life to paper fragments from their nest, or from a foreign conspecific nest or to a neutral condition. Wasps were then transferred to their original nests where recognition abilities were tested. Our results show that wasps do not alter their recognition ability if exposed only to nest material, or in absence of nest material, during the early phase of adult life. It thus appears that the nest paper is not used as a source of recognition cues to be learned in a specific time window, although we discuss possible alternative explanations. Our study provides a novel perspective for the study of the ontogeny of nestmate recognition in Polistes wasps and in other social insects. [ABSTRACT FROM AUTHOR]

Simple Summary: Hornets of the genus Vespa and honey bees are the main characters of a coevolutionary arms race that is made evident by the conspicuous number of reciprocal adaptations evolved by both predator and prey. Through this ancient process predatory hornets and honey bees seem to have reached a fragile balance, which is lost when alien hornet species are accidentally introduced into new geographic areas. This is the case of Asian hornet species, such as Vespa velutina and Vespa mandarinia, which have invaded Europe and North America in recent decades, with a heavy impact on Western honey bees and commercial beekeeping. Here, we provide a comprehensive review of the existing studies that have investigated the interactions between hornets and honey bees throughout the world, highlighting the complexity of the specialized strategies adopted by Asian honey bees and their sympatric predators and the relative lack of coevolution and effective defenses of Western honey bees against invasive hornets. Hornets and honey bees have a long history of coevolution resulting in a plethora of captivating adaptations and counteradaptations between predator and prey. From simple physiological mechanisms to complex behavioral strategies, some Vespa hornets have specialized in hunting honey bees, while the latter have put in place effective defenses to counteract their attack. Both hornets and honey bees have evolved the ability to detect the odors and the pheromones emitted by the other to locate the prey or to spot foraging predators. Hornets often rely on their bigger size, heavily armored body and destructive attacks, while honey bees differentiated collective defense responses finely coordinated to deter or kill the hornet menace. However, when new species of hornets and honey bees come into contact, the absence of coevolution can have a heavy impact on the defenseless bees. The evolutionary arms race between hornets and honey bees provides not only compelling examples of adaptations and counteradaptations between predator and prey, but could also represent a starting point for the development of effective and sustainable strategies to protect honey bees and beekeeping activities and to control invasive alien species of hornets. [ABSTRACT FROM AUTHOR]

Simple Summary: Social insects use cuticular hydrocarbons for chemical recognition and communication. Cuticular hydrocarbons can also be exploited by parasites to their advantage for undermining host recognition systems. The small hive beetle (SHB) is a parasite of honey bee colonies but can also infest nests of other bee species. However, its chemical profile is still not known. For the first time, the present study investigated the SHB chemical profile and compared it with that of its honey bee host. The results show that the SHB has a low chemical profile that is similar to its honey bee host's. However, while honey bees had a clear colony-specific chemical profile, SHBs did not. The generic chemical profile of the SHB is most likely linked to its free-flying behaviour in the field as these parasites are known to switch between host colonies, possibly limiting the acquisition of a colony specific chemical profile. Our findings also suggest that SHBs do not exploit any finely tuned chemical strategy to conceal their presence inside host colonies and probably rely on behavioural adaptations. Cuticular hydrocarbons (CHCs) cover insects' bodies and play important roles in chemical communication, including nestmate recognition, for social insects. To enter colonies of a social host species, parasites may acquire host-specific CHCs or covertly maintain their own CHC profile by lowering its quantity. However, the chemical profile of small hive beetles (SHBs), Aethina tumida, which are parasites of honey bee, Apis mellifera, colonies, and other bee nests, is currently unknown. Here, adults of SHB and honey bee host workers were collected from the same field colonies and their CHC profiles were analysed using GC-MS. The chemical profiles of field-sampled SHBs were also compared with those of host-naive beetles reared in the laboratory. Laboratory-reared SHBs differed in their CHC profiles from field-sampled ones, which showed a more similar, but ten-fold lower, generic host CHC profile compared to host workers. While the data confirm colony-specific CHCs of honey bee workers, the profile of field-collected SHBs was not colony-specific. Adult SHBs often commute between different host colonies, thereby possibly preventing the acquisition of a colony-specific CHC profiles. An ester was exclusive to both groups of SHBs and might constitute an intraspecific recognition cue. Our data suggest that SHBs do not use any finely tuned chemical strategy to conceal their presence inside host colonies and instead probably rely on their hard exoskeleton and defence behaviours. [ABSTRACT FROM AUTHOR]

• Venom volatiles did not differ between female castes in Vespa velutina nigrithorax. • Males were not attracted by venom volatiles. • Sternal gland secretion differed between female castes. • Males were attracted by the females' sternal gland secretion. Due to its huge invasion potential and specialization in honeybee predation, the invasive hornet Vespa velutina nigrithorax represents a high-concern species under both an ecological and economical perspective. In light of the development of specific odorant attractants to be used in sustainable control strategies, we carried out both behavioural assays and chemical analyses to investigate the possibility that, in the invasive population of V. velutina nigrithorax , reproductive females emit volatile pheromones to attract males, as demonstrated in a Chinese non-invasive population. We focused on the secretions produced by sternal and venom glands; because of the volatility and complexity of their composition, both of them could potentially allow an attraction and a species-specific response, decreasing therefore non-target species by-catches. Results of chemical analyses and behavioural assays showed that venom volatiles, although population-specific, are unlikely candidates as male attractants since they do not differ in composition or in quantity between reproductive females and workers and do not attract males. Conversely, sternal gland secretion differs between female castes for the presence of some ketoacids exclusive of gynes already reported as sex pheromones for the non-invasive subspecies V. velutina auraria. Despite such a difference, males are attracted by the sternal gland secretion of both workers and gynes. These results provide a first step to understand the reproductive biology of V. velutina nigrithorax in its invasive range and to develop effective and sustainable management strategies for the species. [ABSTRACT FROM AUTHOR]

In social wasps, female lifespan depends on caste and colony tasks: workers usually live a few weeks while queens as long as 1 year. Polistes dominula paper wasps infected by the strepsipteran parasite Xenos vesparum avoid all colony tasks, cluster on vegetation where parasite dispersal and mating occur, hibernate and infect the next generation of wasp larvae. Here, we compared the survival rate of infected and uninfected wasp workers. Workers' survival was significantly affected by parasite sex: two-third of workers parasitized by a X. vesparum female survived and overwintered like future queens did, while all workers infected by a X. vesparum male died during the summer, like uninfected workers that we used as controls. We measured a set of host and parasite traits possibly associated with the observed lifespan extension. Infected overwintering workers had larger fat bodies than infected workers that died in the summer, but they had similar body size and ovary development. Furthermore, we recorded a positive correlation between parasite and host body sizes. We hypothesize that the manipulation of worker's longevity operated by X. vesparum enhances parasite's fitness: if workers infected by a female overwinter, they can spread infective parasite larvae in the spring like parasitized gynes do, thus contributing to parasite transmission. [ABSTRACT FROM AUTHOR]

The significant risk of disease transmission has selected for effective immune-defense strategies in insect societies. Division of labour, with individuals specialized in immunity-related tasks, strongly contributes to prevent the spread of diseases. A trade-off, however, may exist between phenotypic specialization to increase task efficiency and maintenance of plasticity to cope with variable colony demands. We investigated the extent of phenotypic specialization associated with a specific task by using allogrooming in the honeybee, Apis mellifera, where worker behaviour might lower ectoparasites load. We adopted an integrated approach to characterize the behavioural and physiological phenotype of allogroomers, by analyzing their behavior (both at individual and social network level), their immunocompetence (bacterial clearance tests) and their chemosensory specialization (proteomics of olfactory organs). We found that allogroomers have higher immune capacity compared to control bees, while they do not differ in chemosensory proteomic profiles. Behaviourally, they do not show differences in the tasks performed (other than allogrooming), while they clearly differ in connectivity within the colonial social network, having a higher centrality than control bees. This demonstrates the presence of an immune-specific physiological and social behavioural specialization in individuals involved in a social immunity related task, thus linking individual to social immunity, and it shows how phenotypes may be specialized in the task performed while maintaining an overall plasticity. [ABSTRACT FROM AUTHOR]

Mimicry is one of the most fascinating phenomena in nature 1. Mimicry traits often reflect complex, finely tuned, and sometimes extravagant relationships among species and have evolved to deceive predators or prey. Indeed, mimicry has most often evolved to discourage predation: the 'mimic' exhibits phenotypic convergence towards a non-related 'model' organism which is inedible or harmful, so that a given predator, or 'receiver', will refrain from attacking or ingesting the mimic. Traditionally, mimicry is mainly evident and has been mainly studied in the visual domain. Here, we report experiments that document the first case of interspecific acoustic mimicry in a mammal and demonstrate that the distress calls the greater mouse-eared bat (Myotis myotis) broadcasts when handled imitate sounds of stinging bees or wasps to discourage the bat's avian predators. Ancillotto et al. report a case of interspecific acoustic mimicry in a mammal, and demonstrate that the distress calls of the greater mouse-eared bat imitate sounds of stinging social hymenopterans to discourage the bat's avian predators. [ABSTRACT FROM AUTHOR]

The phenotype of male Hymenoptera and the peculiar role of males has been neglected and greatly understudied, given the spectacular cooperative behavior of female social insects. In social insects there has been considerable progress in understanding the molecular mechanisms behind haplodiploid sex determination but, beyond that, very little is known concerning the neural, endocrine, and genetic correlates of sexual selection in males. An opportunity is being missed: the male phenotype in Hymenoptera is a natural experiment to compare the drives of natural versus sexual selection. In contrast to females, males do not work, they usually display far from the nest to gain mates, compete among rivals in nuptial flights or for a symbolic territory at leks , and engage in direct or ritualized conflicts. By comparing the available data on male paper wasps with studies on other social Hymenoptera, we summarize what we currently know about the physical, hormonal, neural and behavioral traits in a model system appropriate to examine current paradigms on sexual selection. Here we review male behavior in social Hymenoptera beyond sex stereotypes: the subtle role of “drones” in the colony, the lack of armaments and ornaments, the explosive mating crowds, the “endurance” race, the cognitive bases of the “choosy” male and his immune defense. Social insect males are not just simple-minded mating machines, they are shaped, constrained and perhaps trapped by sexual selection. [ABSTRACT FROM AUTHOR]

Paper wasps (Polistes dominula), parasitized by the strepsipteran Xenos vesparum, are castrated and desert the colony to gather on plants where the parasite mates and releases primary larvae, thus completing its lifecycle. One of these plants is the trumpet creeper Campsis radicans: in a previous study the majority of all wasps collected from this plant were parasitized and focused their foraging activity on C. radicans buds. The unexpected prevalence and unusual feeding strategy prompted us to investigate the influence of this plant on wasp behavior and physiology through a multidisciplinary approach. First, in a series of laboratory bioassays, we observed that parasitized wasps spent more time than non-parasitized ones on fresh C. radicans buds, rich of extra-floral nectaries (EFNs), while the same wasps ignored treated buds that lacked nectar drops. Then, we described the structure and ultra-structure of EFNs secreting cells, compatible with the synthesis of phenolic compounds. Subsequently, we analysed extracts from different bud tissues by HPLC-DAD-MS and found that verbascoside was the most abundant bioactive molecule in those tissues rich in EFNs. Finally, we tested the immune-stimulant properties of verbascoside, as the biochemical nature of this compound indicates it might function as an antibacterial and antioxidant. We measured bacterial clearance in wasps, as a proxy for overall immune competence, and observed that it was enhanced after administration of verbascoside—even more so if the wasp was parasitized. We hypothesize that the parasite manipulates wasp behavior to preferentially feed on C. radicans EFNs, since the bioactive properties of verbascoside likely increase host survival and thus the parasite own fitness. [ABSTRACT FROM AUTHOR]

The macroparasite Xenos vesparum affects both the behaviour and the physical traits of its host, the social wasp Polistes dominulus. Female wasps, if parasitized, do not perform any social tasks and desert the colony to gather at specific sites, where the parasite mates; at the end of summer they form prehibernating clusters joined by healthy future queens to overwinter. Parasitized wasps become highly gregarious. In April, healthy wasps leave the aggregations to found new colonies, while parasitized wasps remain in overwintering groups and release parasites to infect wasp larvae only later in the season. We studied the prolonged gregarious behaviour of parasitized wasps and analysed the morphology of parasitized and healthy wasps in aggregations collected over a 7-year period to determine whether the parasite affects host size, wing symmetry, ovarian development and lipid stores. All parasitized wasps were smaller and had undeveloped ovaries and more wing fluctuating asymmetry than unparasitized wasps, irrespective of time of year, parasite load and parasite sex. If infected only by one or two X. vesparum females, the wasps had large fat bodies, which could facilitate their overwintering. In contrast, wasps infected by at least one male parasite had little lipid and died at the end of the summer. Thus, X. vesparum, may play a role in the fate of its host, by exploiting wasps'' tendency to form aggregations outside the colony and by altering its caste system, nutrient allocation, diapause timing and life span to achieve its own reproduction and dispersal. [ABSTRACT FROM AUTHOR]

Social insects are successful animal invaders. Their survival and success, and in some cases also their impact on invaded ecosystem functioning, is often mediated by symbiosis with microorganisms. Here, we report a comprehensive comparative characterization of the gut microbial communities of different castes and developmental stages of the invasive hornet Vespa velutina nigrithorax. The species recently colonized Europe, becoming a high ecological and economic concern, as it threatens pollinator survival and competes with native hornet species. We used targeted meta-genomics to describe the yeasts and bacteria gut communities of individuals of different reproductive phenotypes (workers and future queens), life stages (larvae, newly emerged individuals and adults) and colony non-living samples (nest paper and larval faeces). Bacilli , Gammaproteobacteria , Actinobacteria , Alphaproteobacteria were the most abundant classes of bacteria, and Saccharomycetes , Dothideomycetes , Tremellomycetes and Eurotiomycetes were the most represented yeast classes. We found that the microbial compositions significantly differ across developmental stages and castes, with yeast and bacterial communities switching in frequency and abundance during ontogeny and according to reproductive phenotype. Moreover, the gut microbial communities poorly mirror those found in the nest, suggesting that hornets possess a specific microbial signature. Our results provide the first metagenomic resource of the microbiome of V. velutina in Europe and suggest the importance of considering life stages, reproductive phenotypes and nest influence in order to obtain a comprehensive picture of social insect microbial communities. Unlabelled Image • Vespa velutina is an alien hornet of high ecological and economic concern in Europe. • Its gut microbiome has been characterized through meta-genomics. • We investigated the effect of ontogeny, phenotype and similarity with environment. • Fungal and bacterial community's abundance depends on ontogeny and phenotype. • Microbiome characterization is a springboard for management strategies. [ABSTRACT FROM AUTHOR]

The entomopathogenic fungus Beauveriabassiana is a widely used biopesticide that is considered as an effective alternative to classical agrochemicals. B. bassiana is thought to be safe for pollinators although little is known about its side-effects on pollinators' behaviour and cognition. Here, we focused on honey bees and used the proboscis extension response (PER) protocol to assess whether B. bassiana affects individual sucrose responsiveness, non-associative and associative olfactory learning and memory. Fungus-treated bees displayed an enhanced sucrose responsiveness, which could not be explained by metabolic alterations. Strikingly, exposed bees were twice as inconsistent as controls in response to sucrose, showing PER to lower but not to higher sucrose concentrations. Exposed bees habituated less to sucrose and had a better acquisition performance in the conditioning phase than controls. Further, neither mid- nor long-term memory were affected by the fungus. As sucrose responsiveness is the main determinant of division of foraging labour, these changes might unsettle the numerical ratio between the sub-castes of foragers leading to suboptimal foraging. Although the use of biocontrol strategies should be preferred over chemical pesticides, careful assessment of their side-effects is crucial before claiming that they are safe for pollinators. [ABSTRACT FROM AUTHOR]