Evolution of cuticular hydrocarbons in the hymenoptera : a meta-analysis

Chemical communication is the oldest form of communication, spreading across all organisms of life. In insects, cuticular hydrocarbons (CHC) function as the chemical recognition cues for the recognition of mates, species and nest-mates in social insects. Although much is known about the function of individual hydrocarbons and their biosynthesis, a phylogenetic overview is lacking. Here we review the CHC profiles of 241 species of hymenoptera, one of the largest and important insect orders, including the Symphyta (sawflies), the polyphyletic Parasitica (parasitoid wasps) and the Aculeata (wasps, bees and ants). We investigated whether these five major taxonomic groups differed in the presence and absence of CHC classes and whether the sociality of a species (solitarily vs. social) had an effect on CHC profile complexity. We found that the main CHC classes (i.e., n-alkanes, alkenes and methylalkanes) were all present early in the evolutionary history of the hymenoptera, as evidenced by their presence in ancient Symphyta and primitive Parasitica wasps. Throughout all groups within the Hymenoptera the more complex a CHC the fewer species that produce it, which may reflect the Occam's razor principle that insects’ only biosynthesize the most simple compound that fulfil its needs. Surprisingly there was no difference in the complexity of CHC profiles between social and solitary species, with some of the most complex CHC profiles belonging to the Parasitica. This profile complexity has been maintained in the ants, but some specialisation in biosynthetic pathways has led to a simplification of profiles in the aculeate wasps and bees. The absence of CHC classes in some taxa or species may be due to gene silencing or down-regulation rather than gene loss, as evidenced by sister species having highly divergent CHC profiles, and cannot be predicted by their phylogenetic history. The presence of highly complex CHC profiles prior to the vast radiation of the social hymenoptera indicates a 'spring-loaded' system where the diverse CHC needed for the complex communication systems of social insects, were already present for natural selection to act upon rather than evolve independently. This would greatly aid the multiple evolution of sociality in the Aculeata