Evidence for geographic variation in colony social organization in an obligately social sweat bee, Lasioglossum malachurum Kirby (Hymenoptera; Halictidae)

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Abstract
The expression of altruism and colony eusociality are both a matter of degree in social sweat bees. Even in obligately social species, variation in these traits may be observed across a species' range. Lasioglossum (Evylaeus) malachurum (Kirby) is an obligately eusocial sweat bee found across Europe. In western Europe, L. malachurum exhibits north-south clines of increasing colony size associated with the production of more worker broods, and worker production of males, but societies conform to the model of a classically eusocial hymenopteran insect. A population of L. malachurum studied from 1994 to 1998 at Agios Nikolaos Monemvasias in southern Greece exhibits a startlingly different type of social structure. Dissections of female bees collected while foraging on flowers or from excavations of nests showed that the majority of mid- to late-summer workers are mated and (or) have developing ovaries, indicating that some workers are highly reproductive. Nest excavations indicated that in many or most colonies, the queen has disappeared by midsummer, before ovipositing the final, reproductive brood. In orphan nests, workers become the major reproductives, which suggests that males and gynes in the final brood are the offspring of workers. The very long breeding season in southern Greece may explain why colonies often outlive their queen. The result is the expression of a multivoltine colony cycle and a behavioural switch from eusocial to semisocial colony organization.