Division of labour in social insect colonies relies on behavioural functional differentiation (specialization) of individuals with similar genomes. However, individual behavioural traits do not evolve independently of each other (behavioural syndromes). A prime example is the suite of behavioural differences in honeybee workers that has evolved in response to bidirectional selection on pollen hoarding of honeybee colonies (pollen-hoarding syndrome). More generally, these differences reflect functional differentiation between nectar and pollen foragers. We demonstrate here that this pollen-hoarding syndrome extends to drones. Similar to what has been shown in workers, drones from the high-pollen-hoarding strain had a higher locomotion activity after emergence, and they initiated flight earlier than did males derived from the low-pollen-hoarding strain, with hybrids intermediate. However, these two behavioural traits were unlinked at the individual level. We also found that social environment (the colony) affects the age at which drones initiate flight. The indirect selection responses of male behaviour suggest that male and worker evolution are not independent and may constrain each other's evolution. Furthermore, we identified three distinct peaks in the probability of flight initiation over the course of the experiment and a decreased phenotypic variability in the 'hybrid' males, contrary to quantitative genetic expectations.Social insects owe much of their evolutionary success to the coordinated action of behaviourally specialized members of their colonies (Oster & Wilson 1978; Winston 1987;Hölldobler & Wilson 1990). Division of labour is a hallmark of social evolution (Beshers & Fewell 2001), and among honeybee workers, it is mainly determined by age and genetic effects (Winston 1987). To promote individual specialization, the response thresholds to different task stimuli should vary independently among individuals (Beshers & Fewell 2001). This seems not to be the case: honeybees show well-established behavioural differences among races that extend to multiple behavioural phenotypes, as well as to morphology and life history (Winston 1987). Similar multiple behavioural phenotypes in honeybees are also correlated at the individual level (Page & Erber 2002;Pankiw 2003;Scheiner et al. 2004), indicating their evolution as behavioural syndromes (array of correlated behavioural characteristics: Sih et al. 2004). Linked sets of behavioural traits may have arisen through adaptation in certain contexts (Amdam et al. 2004), but the correlation among behavioural traits may constrain the division of labour in social insects, and behavioural optimization in general (Price & Langen 1992;Sih et al. 2004 One of the best-studied examples of behavioural syndromes in insects is the behavioural differentiation among workers of two strains of honeybees that have been bidirectionally selected over multiple generations for different amounts of pollen stored in their nests . Many different aspects of foraging behaviour Wadd...