How to escape from the host nest: Imperfect chemical mimicry in eucharitid parasitoids and exploitation of the ants’ hygienic behavior

Communication in ants is based to a great extent on chemical compounds. Recognition of intruders is primarily based on cuticular hydrocarbon (CHC) profile matching but is prone to being cheated. Eucharitid wasps are specific parasitoids of the brood of ants; the immature stages are either well integrated within the colony or are protected within the host cocoons, whereas adult wasps at emergence must leave their host nest to reproduce and need to circumvent the ant recognition system to escape unscathed. The behavioral interactions between eucharitid wasps and workers of their host, the Neotropical ant Ectatomma tuberculatum, are characterized. In experimental bioassays, newly emerged parasitoids were not violently aggressed. They remained still and were grabbed by ants upon contact and transported outside the nest; host workers were even observed struggling to reject them. Parasitoids were removed from the nest within five minutes, and most were unharmed, although two wasps (out of 30) were killed during the interaction with the ants. We analyzed the CHCs of the ant and its two parasitoids, Dilocantha lachaudii and Isomerala coronata, and found that although wasps shared all of their compounds with the ants, each wasp species had typical blends and hydrocarbon abundance was also species specific. Furthermore, the wasps had relatively few CHCs compared to E. tuberculatum (22–44% of the host components), and these were present in low amounts. Wasps, only partially mimicking the host CHC profile, were immediately recognized as alien and actively removed from the nest by the ants. Hexane-washed wasps were also transported to the refuse piles, but only after being thoroughly inspected and after most of the workers had initially ignored them. Being recognized as intruder may be to the parasitoids’ advantage, allowing them to quickly leave the natal nest, and therefore enhancing the fitness of these very short lived parasitoids. We suggest that eucharitids take advantage of the hygienic behavior of ants to quickly escape from their host nests.     

Comparative analysis of septic injury-inducible genes in phylogenetically distant model organisms of regeneration and stem cell research, the planarian Schmidtea mediterranea and the cnidarian Hydra vulgaris

Background

Host-parasite interactions are among the most important biotic relationships. Host species should evolve mechanisms to detect their enemies and employ appropriate counterstrategies. Parasites, in turn, should evolve mechanisms to evade detection and thus maximize their success. Females of the European beewolf (Philanthus triangulum, Hymenoptera, Crabronidae) hunt exclusively honeybee workers as food for their progeny. The brood cells containing the paralyzed bees are severely threatened by a highly specialized cuckoo wasp (Hedychrum rutilans, Hymenoptera, Chrysididae). Female cuckoo wasps enter beewolf nests to oviposit on paralyzed bees that are temporarily couched in the nest burrow. The cuckoo wasp larva kills the beewolf larva and feeds on it and the bees. Here, we investigated whether H. rutilans evades detection by its host. Since chemical senses are most important in the dark nest, we hypothesized that the cuckoo wasp might employ chemical camouflage.

Results

Field observations suggest that cuckoo wasps are attacked by beewolves in front of their nest, most probably after being recognized visually. In contrast, beewolves seem not to detect signs of the presence of these parasitoids neither when these had visited the nest nor when directly encountered in the dark nest burrow. In a recognition bioassay in observation cages, beewolf females responded significantly less frequently to filter paper discs treated with a cuticular extract from H. rutilans females, than to filter paper discs treated with an extract from another cuckoo wasp species (Chrysis viridula). The behavior to paper discs treated with a cuticular extract from H. rutilans females did not differ significantly from the behavior towards filter paper discs treated with the solvent only. We hypothesized that cuckoo wasps either mimic the chemistry of their beewolf host or their host's prey. We tested this hypothesis using GC-MS analyses of the cuticles of male and female beewolves, cuckoo wasps, and honeybee workers. Cuticle extracts of Hedychrum nobile (Hymenoptera: Chrysididae) and Cerceris arenaria (Hymenoptera: Crabronidae) were used as outgroups. There was little congruence with regard to cuticular compounds between H. rutilans females and honeybees as well as females of C. arenaria and H. nobile. However, there was a considerable similarity between beewolf females and H. rutilans females. Beewolf females show a striking dimorphism regarding their cuticular hydrocarbons with one morph having (Z)-9-C25:1 and the other morph having (Z)-9-C27:1 as the major component. H. rutilans females were more similar to the morph having (Z)-9-C27:1 as the main component.

Conclusion

We conclude that H. rutilans females closely mimic the composition of cuticular compounds of their host species P. triangulum. The occurrence of isomeric forms of certain compounds on the cuticles of the cuckoo wasps but their absence on beewolf females suggests that cuckoo wasps synthesize the cuticular compounds rather than sequester them from their host. Thus, the behavioral data and the chemical analysis provide evidence that a specialized cuckoo wasp exhibits chemical mimicry of the odor of its host. This probably allows the cuckoo wasp to enter the nest with a reduced risk of being detected by olfaction and without leaving traitorous chemical traces.     

Low level of cuticular hydrocarbons in a parasitoid of a solitary digger wasp and its potential for concealment

Insect cuticular hydrocarbons (CHC) play a role as semiochemicals in many host–parasite systems and chemical mimicry or camouflage is a well-known mechanism of parasites to evade detection by the host. The cuckoo wasp Hedychrum rutilans (Hymenoptera, Chrysididae) is a parasitoid of larvae of the European beewolf Philanthus triangulum (Hymenoptera, Crabronidae). Females chemically mimic the cuticular hydrocarbons of their hosts to avoid detection and countermeasures when entering the host nest for oviposition. Here we report on a possible second mechanism of the chrysidid wasp H. rutilans to evade detection: the amount of CHC/mm² of cuticle is only approximately one-fifth compared to its beewolf host. Furthermore, we show that surprisingly large amounts of CHC of beewolf females can be found on the walls of the underground nest. Potentially, these hydrocarbons might constitute a background odor against which the cuckoo wasps or their chemical traces have to be perceived by the beewolf. The reduction in the amount of CHC of the cuckoo wasps might be equivalent to a dilution of recognition cues, especially against the background odor of the nest walls, and might provide a means to escape detection within the nest due to "chemical insignificance".     

Release from prey preservation behavior via prey switch allowed diversification of cuticular hydrocarbon profiles in digger wasps

The cuticle of insects is covered by a layer of hydrocarbons (CHC), whose original function is the protection from desiccation and pathogens. However, in most insects CHC profiles are species specific. While this variability among species was largely linked to communication and recognition functions, additional selective forces may shape insect CHC profiles. Here, we show that in Philanthinae digger wasps (Crabronidae) the CHC profile coevolved with a peculiar brood‐care strategy. In particular, we found that the behavior to embalm prey stored in the nest with hydrocarbons is adaptive to protect larval food from fungi in those species hunting for Hymenoptera. The prey embalming secretion is identical in composition to the alkene‐dominated CHC profile in these species, suggesting that their profile is adaptively conserved for this purpose. In contrast, prey embalming is not required in those species that switched to Coleoptera as prey. Released from this chemical brood‐care strategy, Coleoptera‐hunting species considerably diversified their CHC profiles. Differential needs to successfully protect prey types used as larval food have thus driven the diversification of CHCs profiles of female Philanthinae wasps. To the best of our knowledge, this is the first evidence of a direct link between selection pressure for food preservation and CHC diversity.     

A mimetic nesting association between a timid social wasp and an aggressive arboreal ant

In French Guiana, the arboreal nests of the swarm-founding social wasp Protopolybia emortualis (Polistinae) are generally found near those of the arboreal dolichoderine ant Dolichoderus bidens. These wasp nests are typically protected by an envelope, which in turn is covered by an additional carton ‘shelter’ with structure resembling the D. bidens nests. A few wasps constantly guard their nest to keep D. bidens workers from approaching. When alarmed by a strong disturbance, the ants invade the host tree foliage whereas the wasps retreat into their nest. Notably, there is no chemical convergence in the cuticular profiles of the wasps and ants sharing a tree. The aggressiveness of D. bidens likely protects the wasps from army ant raids, but the ants do not benefit from the presence of the wasps; therefore, this relationship corresponds to a kind of commensalism.     

Nesting and biology of Jucancistrocerus caspicus (hymenoptera,vespidae, eumeninae)

Females of Jucancistrocerus caspicus nest in dense clay ground on the vertical surface of cliffs. The nests contain 1–9 cells (on average 2.8) and have a linear-branched construction. Females surmount the entrance of the burrow with a curved chimney which has a laced structure. The cells are positioned vertically in the main burrow and obliquely or horizontally in the lateral tunnels; the cells in a row are separated with double partitions. The size of the cells is 7–9 × 4–4.5 mm, the diameter of the nest burrow is 4 mm. The egg is laid before provisioning and is attached to the cell ceiling with a filament. Females hunt for weevil larvae and store 23–33 larvae (on average 27.8) in each cell. The species is univoltine, with prepupae hibernating in their cocoons. The nests are parasitized by the cuckoo wasps Chrysis rutilans which cause 11.5% of brood mortality. Adult wasps are killed by the spiders Pholcus sp. living near the nests.     

Cleptoparasites, social parasites and a common host: Chemical insignificance for visiting host nests, chemical mimicry for living in

Social insect colonies contain attractive resources for many organisms. Cleptoparasites sneak into their nests and steal food resources. Social parasites sneak into their social organisations and exploit them for reproduction. Both cleptoparasites and social parasites overcome the ability of social insects to detect intruders, which is mainly based on chemoreception. Here we compared the chemical strategies of social parasites and cleptoparasites that target the same host and analyse the implication of the results for the understanding of nestmate recognition mechanisms. The social parasitic wasp Polistes atrimandibularis (Hymenoptera: Vespidae), and the cleptoparasitic velvet ant Mutilla europaea (Hymenoptera: Mutillidae), both target the colonies of the paper wasp Polistes biglumis (Hymenoptera: Vespidae). There is no chemical mimicry with hosts in the cuticular chemical profiles of velvet ants and pre-invasion social parasites, but both have lower concentrations of recognition cues (chemical insignificance) and lower proportions of branched alkanes than their hosts. Additionally, they both have larger proportions of alkenes than their hosts. In contrast, post-invasion obligate social parasites have proportions of branched hydrocarbons as large as those of their hosts and their overall cuticular profiles resemble those of their hosts. These results suggest that the chemical strategies for evading host detection vary according to the lifestyles of the parasites. Cleptoparasites and pre-invasion social parasites that sneak into host colonies limit host overaggression by having few recognition cues, whereas post-invasion social parasites that sneak into their host social structure facilitate social integration by chemical mimicry with colony members.     

Antennal responses of an oligolectic bee and its cleptoparasite to plant volatiles

Cleptoparasitic or cuckoo bees lay their eggs in nests of other bees, and the parasitic larvae feed the food that had been provided for the host larvae. Nothing is known about the specific signals used by the cuckoo bees for host nest finding, but previous studies have shown that olfactory cues originating from the host bee alone, or the host bee and the larval provision are essential. Here, I compared by using gas chromatography coupled to electroantennographic detection (GC-EAD) the antennal responses of the oligolectic oil-bee Macropis fulvipes and their cleptoparasite, Epeoloides coecutiens, to dynamic headspace scent samples of Lysimachia punctata, a pollen and oil host of Macropis. Both bee species respond to some scent compounds emitted by L. punctata, and two compounds, which were also found in scent samples collected from a Macropis nest entrance, elicited clear signals in the antennae of both species. These compounds may not only play a role for host plant detection by Macropis, but also for host nest detection by Epeoloides. I hypothesise that oligolectic bees and their cleptoparasites use the same compounds for host plant and host nest detection, respectively.Key words: Macropis fulvipes, Epeoloides coecutiens, Lysimachia punctata, oligolectic oil-bee, floral scent, dynamic headspace, GC-EAD, cuckoo bee, host nest findingBees are the most important animal pollinators worldwide, and guarantee sexual reproduction of many plant species.^1,2 This is especially true for female bees, which collect pollen and mostly nectar for their larvae and frequently visit flowers. For finding and detection of suitable flowers, bees are known to use, besides optical cues,^3,4 especially olfactory signals.^5–8 However, c. 20% of bees do not collect pollen for their larvae by their own, but enter nests of host bees and lay eggs into the broodcells.^1,9 The parasitic larvae subsequently feed the food that had been provided for the host larvae. These so called cuckoo or cleptoparasitic bees can be generalistic, indicating that they use species of several other bee groups as host, whereas others can be highly specialized, laying eggs in cells of only few host species.¹ Until now little is known about the cues used by the cuckoo bees for finding host nests. Nevertheless, Cane¹⁰ and Schindler¹¹ demonstrated that parasitic Nomada bees use primarily visual cues of the nest entrance holes for finding possible nests, and olfactory cues for detection of suitable host nests. The chemical cues used by the cleptoparasites originate from the host bee^10,11 and also pollen,¹⁰ the main larval provision. In most bee species, pollen is mixed together with nectar as larval provision, and both floral resources are known to emit volatiles.^12,13 It is unknown, whether cuckoo bees in search for host nests also use volatiles originating from nectar. While the odours of the host bee used as signal by the cleptoparasites, e.g., cuticiular hydrocarbons and glandular secretions, are often species-specific,¹⁴ the chemical cues from the larval provision may just indicate the presence of pollen in the nest without more specifity. As a consequence cuckoo bees could use species-specific host odours to detect nests of a suitable host, and odours released from the larval provision could indicate to them that broodcells are foraged. However, especially those cuckoo bees with oligolectic hosts foraging pollen only on few closely related plant species,¹ may also use the olfactory signals from host broodcell supplies as more specific cue for host nest detection. Thus the same signal from certain flowers may be used for different informations: for the host bee for host plant and for the cuckoo bee for host nest detection.In this concern I tested oligolectic Macropis (Melittidae, Melittinae) and its specific cuckoo bee, Epeoloides (Apidae, Apinae) by using gas chromatography coupled to electroantennographic detection (GC-EAD) on floral scent of Lysimachia (Myrsinaceae). Macropis is highly specialized on Lysimachia, because it is not only collecting pollen from plants of this genus, but also floral oil. Both floral products are the only provision for the larvae.^1,15 Recently, we have shown that the oil bee Macropis is strongly attracted to floral scent of its oil host Lysimachia though the compounds used for host plant finding are still unknown.⁷ Macropis is the only host of Epeoloides, and larvae of this cleptoparasite only feed on the Lysimachia pollen-oil mixture provided for the larvae of Macropis. Worldwide, there are only 2 species of this genus, one in North America and the other in Europe/Asia.^1,16,17 I hypothesized that both bee species respond to specific Lysimachia compounds, which may be used for host plant as well as host nest detection.The measurements with M. fulvipes (F.) and E. coecutiens (F.) antennae demonstrate that both bees, host as well as cuckoo bee, respond to some scent compounds emitted by inflorescences of Lysimachia punctata L. (Fig. 1), a plant being an important pollen and oil source for M. fulvipes. Macropis responded to much more Lysimachia compounds compared to the cuckoo bee, however, two compounds elicited clear signals in the antennae of both bee species: the benzenoid 1-hydroxy-1-phenyl-2-propanone, and the fatty acid derivative 2-tridecanone. Interestingly, both compounds are also emitted from the floral oil of this plant,⁷ and both compounds were also detected in scent samples collected by dynamic headspace in the entrance of a Macropis nest (Dötterl, unpublished data). Therefore, an Epeoloides female being in search for a host nest can detect volatiles emitted from the provision of the host bee at the entrance of a bee nest, and may use these specific compounds for detection of a Macropis nest provisioned with Lysimachia pollen and oil.Open in a separate windowFigure 1Coupled gas chromatographic and electroantennographic detection of a Lysimachia punctata headspace scent sample using antennae of a female oligolectic Macropis fulvipes and a female cleptoparasitic Epeoloides coecutiens bee. (1) 1-hydroxy-1-phenyl-2-propanone, (2) 2-tridecanone.Present results show that an oligolectic oil-bee as well as its cleptoparasite detects volatiles originating from the host plant of the pollen collecting bee, and that oligolectic bees as well as their cuckoo bees may use the same specific signals for host plant and host nest finding, respectively. Biotests are now needed to test this hypothesis.     

Cuticular hydrocarbon dynamics in young adult Polistes dominulus (Hymenoptera: Vespidae) and the role of linear hydrocarbons in nestmate recognition systems

In social insects, cuticular hydrocarbons (CHCs) play an important role in nestmate discrimination processes, but young individuals are usually not discriminated. We studied CHC changes in young workers of the social wasp Polistes dominulus. A quantitative estimation demonstrated that total quantities of CHCs increased after emergence, with branched alkanes increasing drastically when compared with other classes of hydrocarbons. The relative quantity of longer-chain compounds increased with respect to shorter ones; unsaturated compounds decreased. These changes might reduce the capacity of the cuticle to acquire compounds of environmental origin. We then tested whether individuals acquire hydrocarbons from the environment, and whether this capability equally characterises newly emerged and mature wasps. We exposed wasps of two age classes (adults younger or older than 24 h) to four linear hydrocarbons in turn, and observed how nestmates reacted to their re-introduction into the natal colony. Exposed young wasps elicited significantly more aggressive responses than control sisters; but treated wasps older than 24 h were generally accepted by nestmates. Chemical assays showed that exposed young wasps readily absorbed hydrocarbons; older ones did not incorporate hydrocarbons, suggesting that the chemical profiles of mature wasps are less prone to chemical shifts than those of newly emerged wasps.     

Premating isolation is determined by larval rearing substrates in cactophilic Drosophila mojavensis. IX. Host plant and population specific epicuticular hydrocarbon expression influences mate choice and sexual selection

Sexual signals in cactophilic Drosophila mojavensis include cuticular hydrocarbons (CHCs), contact pheromones that mediate female discrimination of males during courtship. CHCs, along with male courtship songs, cause premating isolation between diverged populations, and are influenced by genotype × environment interactions caused by different host cacti. CHC profiles of mated and unmated adult flies from a Baja California and a mainland Mexico population of D. mojavensis reared on two host cacti were assayed to test the hypothesis that male CHCs mediate within‐population female discrimination of males. In multiple choice courtship trials, mated and unmated males differed in CHC profiles, indicating that females prefer males with particular blends of CHCs. Mated and unmated females significantly differed in CHC profiles as well. Adults in the choice trials had CHC profiles that were significantly different from those in pair‐mated adults from no‐choice trials revealing an influence of sexual selection. Females preferred different male CHC blends in each population, but the influence of host cactus on CHC variation was significant only in the mainland population indicating population‐specific plasticity in CHCs. Different groups of CHCs mediated female choice‐based sexual selection in each population suggesting that geographical and ecological divergence has the potential to promote divergence in mate communication systems.     

Phylogenetic analysis of the mitochondrial genes LSU rRNA and COI suggests early adaptive differentiation of anal teeth in chrysidine cuckoo wasps (Hymenoptera: Chrysididae)

Cuckoo wasps are a morphologically diverse group of Hymenoptera with parasitoid or cleptoparasitic life histories. In the present paper, we explore the phylogenetic signal in fragments of the mitochondrial genes LSU rRNA and COI to resolve the group's phylogeny. We analyzed sequence data of 33 species representing the taxa Cleptinae, Elampini, Parnopini, and Chrysidini. Most of the currently recognized relationships of major cuckoo wasp lineages are supported by the molecular data. A key difference concerns the phylogenetic position of the Euchroeus (=Brugmoia) group within the tribe Chrysidini. It seems likely that an erroneous interpretation of morphological characters has led to inappropriate rooting of that tribe. We suggest that species of the Euchroeus group be interpreted as forming the stem group of the Chrysidini and that the remaining genera of that tribe be united in a subordinated taxon. Our results imply that the evolution of anal dentition, of significance for breaking into sealed host nests otherwise not accessible to cuckoo wasps, already happened at the base of the Chrysidini and that an even number of anal teeth arose prior to an odd number.     

Insights into the structure of plant-insect communities: Specialism and generalism in a regional set of non-pollinating fig wasp communities

Insects show a multitude of symbiotic interactions that may vary in degree of specialization and structure. Gall-inducing insects and their parasitoids are thought to be relatively specialized organisms, but despite their ecological importance, the organization and structure of the interactions they establish with their hosts has seldom been investigated in tropical communities. Non-pollinating fig wasps (NPFW) are particularly interesting organisms for the study of ecological networks because most species strictly develop their offspring within fig inflorescences, and show a multitude of life history strategies. They can be gall-makers, cleptoparasites or parasitoids of pollinating or of other non-pollinating fig wasps. Here we analysed a set of non-pollinating fig wasp communities associated with six species of Ficus section Americanae over a wide area. This allowed us to investigate patterns of specialization in a diverse community composed of monophagous and polyphagous species. We observed that most NPFW species were cleptoparasites and parasitoids, colonizing figs several days after oviposition by pollinators. Most species that occurred in more than one host were much more abundant in a single preferential host, suggesting specialization. The food web established between wasps and figs shows structural properties that are typical of specific antagonistic relationships, especially of endophagous insect networks. Two species that occurred in all available hosts were highly abundant in the network, suggesting that in some cases generalized species can be more competitive than strict specialists. The Neotropical and, to a lesser extent, Afrotropical NPFW communities seem to be more generalized than other NPFW communities. However, evidence of host sharing in the Old World is quite limited, since most studies have focused on particular taxonomic groups (genera) of wasps instead of sampling the whole NPFW community. Moreover, the lack of quantitative information in previous studies prevents us from detecting patterns of host preferences in polyphagous species.     

Strongly biased sex ratio in cuckoo wasp Chrysura hirsuta (Hymenoptera: Chrysididae), a parasitoid of the mason bee Osmia orientalis

We examined the rate of parasitism and sex ratio of the cuckoo wasp Chrysura hirsuta (Gerstaecker) (Hymenoptera: Chrysididae) that emerged from nests of the mason bee Osmia orientalis Benoist (Hymenoptera: Megachilidae) in Nara, Japan. Nests of O. orientalis were found in empty shells of two snail species, Satsuma japonica (Pfeiffer) and Euhadra amaliae (Kobelt). The percentage of parasitism by cuckoo wasps per all collected cocoons tended to be high (20–50%) even though interannual variation and the average number of cocoons per nest did not differ across snail shell species within each year. Our results from three years of observation, combined with previous reports, showed that the adult sex ratio of C. hirsuta was strongly female‐biased, which suggests that the species reproduces by thelytokous parthenogenesis.     

Cuticular hydrocarbon profiles of codling moth larvae,Cydia pomonella (Lepidoptera: Tortricidae), reflect those of their host plant species

The cuticular hydrocarbons (CHCs) of codling moth larvae collected in the field from their host plant species, apple and walnut, were analyzed and compared with the CHCs of fruits from these two phylogenetically distant hosts. The CHC profiles of the larvae consisted solely of n‐alkanes (C₂₃–C₃₁) and differed quantitatively between host populations. Amounts of the CHCs from the walnut‐collected larvae were shifted towards longer‐chain alkanes compared to those from apple‐collected larvae. A similar shift was observed for the CHC profiles of walnut and apple fruits. Analysis of the CHCs of larvae reared on artificial diet, in comparison with hydrocarbons from the diet, confirmed that larval CHCs scarcely reflect hydrocarbons from the food source. This finding indicates that the larval hydrocarbons must be biosynthesized to a large degree by the insect, rather than being gained directly from its diet. Hence, codling moth populations from apple and walnut each synthesize their own CHC profiles, which largely resemble those of their respective host plant, yielding a potential tool of chemical camouflage from certain natural antagonists of the larvae. The findings of the present study, together with recent molecular population analyses, provides evidence for a process that might ultimately lead to sympatric speciation of this herbivore species. © 2010 The Linnean Society of London, Biological Journal of the Linnean Society, 2010, 101 , 376–384.     

Nestling cuckoos,Cuculus canorus,exploit hosts with begging calls that mimic a brood

Nestling cuckoos, Cuculus canorus, eject host eggs or young from the nest and are then raised alone by the hosts. Using reed warblers, Acrocephalus scirpaceus, as hosts, we investigated how the single cuckoo chick can command the same provisioning rate as a whole brood of host young. Large size alone is not sufficient to stimulate adequate provisioning because single blackbird, Turdus merula, or song thrush, T. philomelos, chicks of the same mass as a cuckoo were fed at a lower rate. Our experiments show that the key stimulus is the cuckoo chick''s rapid begging call (''si, si, si, si ...''), which sounds remarkably like a whole brood of host chicks, and which it matched in calling rate. When single blackbird or song thrush chicks were accompanied by loudspeakers that broadcast either cuckoo begging calls or calls of a brood of reed warblers, the hosts increased their provisioning rate to that for a cuckoo chick. We suggest that the cuckoo needs vocal trickery to stimulate adequate care to compensate for the fact that it presents a visual stimulus of just one gape.     

Palaeocene origin of the Neotropical lineage of cleptoparasitic bees Ericrocidini‐Rhathymini (Hymenoptera,Apidae)

Cleptoparasitic bees abandoned pollen‐collecting for their offspring and lay their eggs in other bees' provisioned nests. Also known as cuckoo bees, they belong to several lineages, but are especially diverse in Apinae. We focused on a lineage of apine cleptoparasitic bees, the clade Ericrocidini + Rhathymini, which attack nests of oil‐collecting bees. We sequenced five genes for 20 species in all genera of this clade plus a large outgroup to reconstruct the phylogeny and estimate divergence times. We confirmed the monophyly of the clade Ericrocidini + Rhathymini and its position inside the ericrocidine line together with the tribes Protepeolini, Isepeolini, Osirini and Coelioxoidini. Our results corroborated the current taxonomic classification. Ericrocis is sister to all Ericrocidini and the position of Acanthopus and the most diverse genus Mesoplia were inconclusive. Ericrocidini + Rhathymini diverged from other apine cleptoparasitic lineages 74 Ma in the Cretaceous and from each other in the Palaeocene at 61 Ma. Considering the robust molecular evidence of their sister relationships, the striking differences in the morphology of first‐instar larvae of the two groups may represent adaptations to the nesting biology of their hosts. As other cleptoparasites in the ericrocidine line, Ericrocidini and Rhathymini possess larvae which are adapted to kill the immature host and to feed on floral oil provided by the host female. The evolution of host specialization in the line Ericrocidini + Rhathymini retroceded to the Eocene when they differentiated synchronously with their hosts, Centris and Epicharis .     

Ityphilus krausi N.Sp., a New Ballophilid Centipede from Peru (Chilopoda: Geophilomorpha: Ballophilidae)

The Amazon rainforest is one of the planet’s biodiversity hotspots, hosting a rich orchid bee fauna. The phoretic cleptoparasites of this bee fauna are largely unknown. We report for the first time the host–cleptoparasite interaction between Eulaema mocsaryi (Friese) (Hymenoptera: Apidae: Euglossini) and the first instar larva (triungulin) of a Tetraonycini meloid beetle. We review the host–cleptoparasite interactions of Tetraonycini with Apid bees in South America and discuss the ecological needs of the cleptoparasite.     

Convergence of chemical mimicry in a guild of aphid predators

Abstract.  1. A variety of insects prey on honeydew-producing Homoptera and many do so even in the presence of ants that tend, and endeavour to protect, these trophobionts from natural enemies. Few studies have explored the semiochemical mechanisms by which these predators circumvent attack by otherwise aggressive ants.
2. Ants use specific mixtures of cuticular hydrocarbons (CHCs) as recognition labels, but this simple mechanism is frequently circumvented by nest parasites that engage in 'chemical mimicry' of their host ants by producing or acquiring a critical suite of these CHCs.
3. Analysis of the CHCs from the North American woolly alder aphid, Prociphilus tessellatus (Homoptera: Aphididae), their tending ants, and aphid predators from three insect orders, Feniseca tarquinius (Lepidoptera: Lycaenidae), Chrysopa slossonae (Neuroptera: Chrysopidae), and Syrphus ribesii (Diptera: Syrphidae), showed that while the CHC profile of each predatory species was distinct, each was chemically more similar to the aphids than to either tending ant species. Further, the CHCs of each predator species were a subset of the compounds found in the aphids' profile.
4. These results implicate CHCs as a recognition cue used by ants to discriminate trophobionts from potential prey and a probable mechanism by which trophobiont predators circumvent detection by aphids and their tending ants.
5. Although several features of the aphids' CHC profile are shared among the chemically mimetic taxa, variation in the precision of mimicry among the members of this predatory guild demonstrates that a chemical mimic need not replicate every feature of its model.     

Female choice for male cuticular hydrocarbon profile in decorated crickets is not based on similarity to their own profile

Indirect genetic benefits derived from female mate choice comprise additive (good genes) and nonadditive genetic benefits (genetic compatibility). Although good genes can be revealed by condition‐dependent display traits, the mechanism by which compatibility alleles are detected is unclear because evaluation of the genetic similarity of a prospective mate requires the female to assess the genotype of the male and compare it to her own. Cuticular hydrocarbons (CHCs), lipids coating the exoskeleton of most insects, influence female mate choice in a number of species and offer a way for females to assess genetic similarity of prospective mates. Here, we determine whether female mate choice in decorated crickets is based on male CHCs and whether it is influenced by females' own CHC profiles. We used multivariate selection analysis to estimate the strength and form of selection acting on male CHCs through female mate choice, and employed different measures of multivariate dissimilarity to determine whether a female's preference for male CHCs is based on similarity to her own CHC profile. Female mating preferences were significantly influenced by CHC profiles of males. Male CHC attractiveness was not, however, contingent on the CHC profile of the choosing female, as certain male CHC phenotypes were equally attractive to most females, evidenced by significant linear and stabilizing selection gradients. These results suggest that additive genetic benefits, rather than nonadditive genetic benefits, accrue to female mate choice, in support of earlier work showing that CHC expression of males, but not females, is condition dependent.     

Born in an Alien Nest : How Do Social Parasite Male Offspring Escape from Host Aggression?

Social parasites exploit the colony resources of social insects. Some of them exploit the host colony as a food resource or as a shelter whereas other species also exploit the brood care behavior of their social host. Some of these species have even lost the worker caste and rely completely on the host''s worker force to rear their offspring. To avoid host defenses and bypass their recognition code, these social parasites have developed several sophisticated chemical infiltration strategies. These infiltration strategies have been highly studied in several hymenopterans. Once a social parasite has successfully entered a host nest and integrated its social system, its emerging offspring still face the same challenge of avoiding host recognition. However, the strategy used by the offspring to survive within the host nest without being killed is still poorly documented. In cuckoo bumblebees, the parasite males completely lack the morphological and chemical adaptations to social parasitism that the females possess. Moreover, young parasite males exhibit an early production of species-specific cephalic secretions, used as sexual pheromones. Host workers might thus be able to recognize them. Here we used a bumblebee host-social parasite system to test the hypothesis that social parasite male offspring exhibit a chemical defense strategy to escape from host aggression during their intranidal life. Using behavioral assays, we showed that extracts from the heads of young cuckoo bumblebee males contain a repellent odor that prevents parasite males from being attacked by host workers. We also show that social parasitism reduces host worker aggressiveness and helps parasite offspring acceptance.