-Social insect colonies provide a stable and safe environment for their 34 members. Despite colonies been heavily guarded, parasites have evolved numerous 35 strategies to invade and inhabit these hostile places. Two common strategies are chemical 36 mimicry via biosynthesis of the hosts' odour or chemical camouflage were compounds 37 are acquired straight from the host. The ectoparasitic mite Varroa destructor feeds on the 38 heamolymph of its honeybee host Apis mellifera and uses chemical mimicry to remain 39 undetected as it lives on the adult host during its phoretic phase or while reproducing on 40 the honeybee brood.. During the mite life cycle it switches between host adults and 41 brood, which requires it to adjust its profile to mimic the very different odours of 42 honeybee brood and adults. In a series of transfer experiments using adult bees and 43 pupae, we tested whether V. destructor does this by synthesising compounds or using 44 chemical camouflage. We show that V. destructor required direct access to the host 45 cuticle to mimic its odour and was unable to synthesise host-specific compounds itself. destructor was adjusted within three to nine hours after switching hosts, demonstrating 49 that passive camouflage is a highly efficient, fast and flexible way for the mite's to adapt 50 to a new host's profile when moving between different host life stages, or host colonies.

Phenotypic variation arises from interactions between genotype and environment, although how variation is produced and then maintained remains unclear. The discovery of the nest-mate recognition system in Formica exsecta ants has allowed phenotypic variation in chemical profiles to be quantified across a natural population of 83 colonies. We investigated if this variation was correlated or not with intrinsic (genetic relatedness), extrinsic (location, light, temperature), or social (queen number) factors. (Z)-9-Alkenes and n-alkanes showed different patterns of variance: island (location) explained only 0.2 % of the variation in (Z)-9-alkenes, but 21–29 % in n-alkanes, whereas colony of origin explained 96 % and 45–49 % of the variation in (Z)-9-alkenes and n-alkanes, respectively. By contrast, within-colony variance of (Z)-9-alkenes was 4 %, and 23–34 % in n-alkanes, supporting the function of the former as recognition cues. (Z)-9-Alkene and n-alkane profiles were correlated with the genetic distance between colonies. Only n-alkane profiles diverged with increasing spatial distance. Sampling year explained a small (5 %), but significant, amount of the variation in the (Z)-9-alkenes, but there was no consistent directional trend. Polygynous colonies and populous monogynous colonies were dominated by a rich C23:1 profile. We found no associations between worker size, mound exposure, or humidity, although effect sizes for the latter two factors were considerable. The results support the conjecture that genetic factors are the most likely source of between-colony variation in cuticular hydrocarbons.Electronic supplementary materialThe online version of this article (doi:10.1007/s10886-013-0366-0) contains supplementary material, which is available to authorized users.

Chemical recognition cues are used to discriminate among species, con-specifics, and potentially between patrilines in social insect colonies. There is an ongoing debate about the possible persistence of patriline cues despite evidence for the mixing of colony odors via a “gestalt” mechanism in social insects, because patriline recognition could lead to nepotism. We analyzed the variation in recognition cues (cuticular hydrocarbons) with different mating frequencies or queen numbers in 688 Formica exsecta ants from 76 colonies. We found no increase in the profile variance as genetic diversity increased, indicating that patriline effects were absent or possibly obscured by a gestalt mechanism. We then demonstrated that an isolated individual's profile changed considerably relative to their colony profile, before stabilizing after 5 days. We used these isolated individuals to eliminate the masking effects of the gestalt mechanism, and we detected a weak but statistically significant patriline effect in isolated adult workers and also in newly emerged callow workers. Thus, our evidence suggests that genetic variation in the cuticular hydrocarbon profile of F. exsecta ants (n-alkanes and alkenes) resulted in differences among patrilines, but they were obscured in the colony environment, thereby avoiding costly nepotistic behaviors.

Among social insects, maintaining a distinct colony profile allows individuals to distinguish easily between nest mates and non-nest mates. In ants, colony-specific profiles can be encoded within their cuticular hydrocarbons, and these are influenced by both environmental and genetic factors. Using nine monogynous Formica exsecta ant colonies, we studied the stability of their colony-specific profiles at eight time points over a 4-year period. We found no significant directional change in any colony profile, suggesting that genetic factors are maintaining this stability. However, there were significant short-term effects of season that affected all colony profiles in the same direction. Despite these temporal changes, no significant change in the profile variation within colonies was detected: each colony's profile responded in similar manner between seasons, with nest mates maintaining closely similar profiles, distinct from other colonies. These findings imply that genetic factors may help maintain the long-term stability of colony profile, but environmental factors can influence the profiles over shorter time periods. However, environmental factors do not contribute significantly to the maintenance of diversity among colonies, since all colonies were affected in a similar way.

Our understanding of the role of cuticular hydrocarbons (CHC) in recognition is based largely on temperate ant species and honey bees. The stingless bees remain relatively poorly studied, despite being the largest group of eusocial bees, comprising more than 400 species in some 60 genera. The Meliponini and Apini diverged between 80–130 Myr B.P. so the evolutionary trajectories that shaped the chemical communication systems in ants, honeybees and stingless bees may be very different. The aim of this study was to study if a unique species CHC signal existed in Neotropical stingless bees, as has been shown for many temperate species, and what compounds are involved. This was achieved by collecting CHC data from 24 colonies belonging to six species of Melipona from North-Eastern Brazil and comparing the results with previously published CHC studies on Melipona. We found that each of the eleven Melipona species studied so far each produced a unique species CHC signal based around their alkene isomer production. A remarkable number of alkene isomers, up to 25 in M. asilvai, indicated the diversification of alkene positional isomers among the stingless bees. The only other group to have really diversified in alkene isomer production are the primitively eusocial Bumblebees (Bombus spp), which are the sister group of the stingless bees. Furthermore, among the eleven Neotropical Melipona species we could detect no effect of the environment on the proportion of alkane production as has been suggested for some other species.Electronic supplementary materialThe online version of this article (10.1007/s10886-017-0901-5) contains supplementary material, which is available to authorized users.