1. Invasive alien species are a major threat to biodiversity. In addition to predation and parasitism, native species might suffer from competition when invasive alien species occupy a similar ecological niche.2. This study focused on the potential interspecific interaction between two hornets: the Asian yellow-legged hornet, Vespa velutina, a high-concern invasive alien species recently arrived in Europe; and the native European hornet, Vespa crabro. The two species share a similar ecological niche and V. velutina is rapidly expanding across Europe, which suggests that V. crabro might suffer from competition.3. Under laboratory-controlled conditions, two life-history traits that might cause the two species to compete were investigated: (i) the ability of workers to find food sources and their flexibility in exploiting them (through individual food item choice tests and exploration assays); and (ii) the worker resistance to pathogens (through immune challenge tests).4. The results show that trophic preference of both species highly overlaps, with a marked dietary preference for honeybees compared with other insect prey and non-prey protein items. No differences were observed in the exploratory behaviour of both species. Finally, constitutive antibacterial activity was greater in workers of the native species than in workers of the invasive hornet.5. This laboratory study provides a first assessment under controlled conditions of the factors affecting competition between workers of two hornet species and proposes a framework to assess, in wild contexts, the magnitude of the competition and the impact of the introduced V. velutina on the native V. crabro.

Keywords:animal communication badge of status conventional signal dominance hierarchy hover wasp Liostenogaster flavolineata Liostenogaster vechti quality signal recognition system social behaviour Both recognition and conventional signals are widespread in the animal kingdom. Chemical communication plays a major role in invertebrates, and especially in social insects. In the last decade, observational and experimental evidence has shown the existence of visual quality signals and individual recognition cues in Polistes paper wasps, meaning that visual communication might also be common in insect societies. Here we show that two species of facultatively eusocial hover wasps (Vespidae, Stenogastrinae) use the visual channel for social communication. By combining morphoanatomical measurements and behavioural assays, we found that the size of the dark facial markings was related to reproductive status and dominance in colonies of Liostenogaster vechti, thus representing a badge of status. By contrast, no correlation between facial coloration and reproductive status was found in Liostenogaster flavolineata, which instead used facial markings as familiar visual recognition cues. Our results reveal that visual communication in social insects might be more widespread than previously thought and has evolved independently in distinct wasp taxa facing similar selection pressures. Ó

Honeybee colonies are under the threat of many stressors, biotic and abiotic factors that strongly affect their survival. Recently, great attention has been directed at chemical pesticides, including their effects at sub-lethal doses on bee behaviour and colony success; whereas the potential side effects of natural biocides largely used in agriculture, such as entomopathogenic fungi, have received only marginal attention. Here, we report the impact of the fungus Beauveria bassiana on honeybee nestmate recognition ability, a crucial feature at the basis of colony integrity. We performed both behavioural assays by recording bee guards’ response towards foragers (nestmate or non-nestmate) either exposed to B. bassiana or unexposed presented at the hive entrance, and GC-MS analyses of the cuticular hydrocarbons (CHCs) of fungus-exposed versus unexposed bees. Our results demonstrated that exposed bees have altered cuticular hydrocarbons and are more easily accepted into foreign colonies than controls. Since CHCs are the main recognition cues in social insects, changes in their composition appear to affect nestmate recognition ability at the colony level. The acceptance of chemically unrecognizable fungus-exposed foragers could therefore favour forager drift and disease spread across colonies.

Social insects have evolved sophisticated recognition systems enabling them to accept nest-mates but reject alien conspecifics. In the social wasp, Liostenogaster flavolineata (Stenogastrinae), individuals differ in their cuticular hydrocarbon profiles according to colony membership; each female also possesses a unique (visual) facial pattern. This species represents a unique model to understand how vision and olfaction are integrated and the extent to which wasps prioritize one channel over the other to discriminate aliens and nest-mates. Liostenogaster flavolineata females are able to discriminate between alien and nest-mate females using facial patterns or chemical cues in isolation. However, the two sensory modalities are not equally efficient in the discrimination of 'friend' from 'foe'. Visual cues induce an increased number of erroneous attacks on nest-mates (false alarms), but such attacks are quickly aborted and never result in serious injury. Odour cues, presented in isolation, result in an increased number of misses: erroneous acceptances of outsiders. Interestingly, wasps take the relative efficiencies of the two sensory modalities into account when making rapid decisions about colony membership of an individual: chemical profiles are entirely ignored when the visual and chemical stimuli are presented together. Thus, wasps adopt a strategy to 'err on the safe side' by memorizing individual faces to recognize colony members, and disregarding odour cues to minimize the risk of intrusion from colony outsiders.

The two paper wasps, Polistes dominula and Polistes gallicus, are related species with strongly differing distribution ranges. We investigated thermal tolerance traits (critical thermal limits and metabolic response to temperature) to gain knowledge about physiological adaptations to their local climate conditions and to get evidence for the reasons of P. dominula’s successful dispersion. Body and ambient temperature measurements at the nests revealed behavioural adaptations to microclimate. The species differed clearly in critical thermal minimum (P. dominula −1.4 °C, P. gallicus −0.4 °C), but not significantly in critical thermal maximum of activity (P. dominula 47.1 °C, P. gallicus 47.6 °C). The metabolic response did not reveal clear adaptations to climate conditions. At low and high temperatures, the metabolic rate of P. dominula was higher, and at intermediate temperatures, we determined higher values in P. gallicus. However, the species exhibited remarkably differing thermoregulatory behaviour at the nest. On average, P. gallicus tolerated a thoracic temperature up to ~41 °C, whereas P. dominula already tried at ~37 °C to keep the thorax below ambient temperature. We suggest this to be an adaptation to the higher mean ambient temperature we measured at the nest during a breeding season. Although we determined for P. dominula a 0.5 °C larger thermal tolerance range, we do not presume this parameter to be solely responsible for the successful distribution of P. dominula. Additional factors, such as the thermal tolerance of the queens could limit the overwintering success of P. gallicus in a harsher climate.

The occurrence of a pre-imaginal caste determination represents a sort of "point of no return" to eusociality. In some social insect taxa, including Polistes species, the occurrence of distinct queen and worker castes is still debated. Before this report, no clear morphological diVerences between reproductive and non-reproductive individuals were known. Here we show that several pre-imaginal morphological diVerences of the Van der Vecht organ occur between foundresses (queens) and workers. Geometric morphometrics revealed that queens are characterized by shape deformations of this organ, which is responsible for a typical allometric growth of the secretory area. This organ is predicted to be larger in foundresses compared to workers because its secretion is involved in defense against ants, in nestmate recognition and in preventing workers from challenging for direct reproduction. The results presented here indicate the existence of an incipient morphological caste determination Polistes gallicus and suggest that this species may have passed the "point of no return" for eusociality.

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In the paper wasp Polistes dominula, cuticular hydrocarbons play a critical role to acquire information regarding conspecific individuals. However, the relationship between cuticular hydrocarbons, health status, and male sexually selected traits is poorly investigated. In this study, we characterized the cuticular hydrocarbon profile of adult male and female wasps, infected or not by the strepsipteran endoparasite Xenos vesparum, to assess whether the chemical signature provides information about sex and health status (parasite infection). Moreover, we tested whether the chemical profile reflects male quality as measured via morphological and behavioural (sexually selected) traits at leks. Our results showed that males and females had similar total amount of CHCs, quantitatively different profiles and, to a lesser extent, sex-specific chemical compounds. Cuticular profiles were influenced by the strepsipteran infection, and the effect was stronger in females (the primary host) than in males, according to the physiological castration of female but not of male hosts. Regarding territorial and non-territorial males, no significant difference emerged in their chemical profiles. Furthermore, sex-dimorphic visual signals (size, shape, and asymmetry of abdominal yellow spots) were related to the behavioural displays of territorial males. We hypothesize that cuticular hydrocarbons are potential multi-role cues to assess sex and health status in male and female wasps, in synergy with visual signals and territorial performance in signaling male quality.