Sources of variation in cuticular hydrocarbons in the ant Formica exsecta.

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.