Aggressive and Docile Colony Defence Patterns in Apis mellifera. A Retreater-Releaser Concept

Colony defence in Apis mellifera involves a variety of traits ranging from 'aggressive' (e.g. entrance guarding, recruitment of flying guards) to 'docile' (e.g. retreating into the nest) expression. We tested 11 colonies of three subspecies (capensis, scutellata, carnica) regarding their defensiveness. Each colony was selected as reportedly 'aggressive', 'intermediate' or 'docile' and consisted of about 10,000 bees. We applied three stimulation regimes (mechanical disturbance, exposure to alarm pheromones, and the combination of both) and measured their behaviours by tracking the rates of outflying bees at the entrance sites of the test hives. We provided evidence that for mechanical disturbances the test colonies resolved into two response types, if the 'immediate' defence response, assessed in the first minute of stimulation, was taken as a function of foraging: 'releaser' colonies allocated flying guards, 'retreater' colonies reduced the outside-hive activities. This division was observed irrespective of the subspecies membership and maintained in even roughly changing environmental conditions. However, if pheromone and mechanical stimulation were combined, the variety of colony defensiveness restricted to two further types irrespective of the subspecies membership: six of nine colonies degraded their rate of flying defenders with increasing foraging level, three of the colonies extended their 'aggressiveness' by increasing the defender rate with the foraging level. Such 'super-aggressive' colonies obviously are able to allocate two separate recruitment pools for foragers and flying defenders.