Do hornets have zombie workers?

AbstractColonies of the European hornet,Vespa crabro, are typically founded by a single queen mated to a single male.From the resulting colony relatedness pattern we predicted strong worker–queen conflict over male production where both the workers and the queen attempt to produce the colony’s males. To test for this conflict, male production was studied in 15 hornet nests using a combination of DNA microsatellite analysis (282 males), worker ovary dissections (500 workers from eight nests) and 50 h of observation (four nests). In contrast to our prediction, the data show that hornet males are queens’ sons, that workers never attempt to lay eggs, rarely have activated ovaries, and that there is no direct aggression between the queen and the workers. This contrasts with other data for vespine wasps, which support relatedness predictions.Dolichovespula arenariahas the same kin structure asV. crabroand workers produce males in many colonies. The similarity between these two species makes it difficult to explain why workers do not reproduce inV. crabro. Self‐restraint is expected if worker reproduction significantly reduces colony productivity but there is no obvious reason why this should be important toV. crabrobut not toD. arenaria. Alternatively, queen control may be important. The absence of expressed queen–worker conflict rules out physical control. Indirect pheromonal control is a possibility and is supported by the occurrence of royal courts and queen pheromone inVespabut notDolichovespula. Pheromonal queen control is considered evolutionarily unstable, but could result from a queen–worker arms race over reproductive control in which the queen is ahead. The genetic data also revealed diploid males in one colony, the first example in the vespine wasps, and two colonies with double matrilines, suggesting that occasional usurpation by spring queens occurs.