Abstract
Shimmering is a collective defence behaviour in Giant honeybees (Apis dorsata) whereby individual bees flip their abdomen upwards, producing Mexican wave-like patterns on the nest surface. Bucket bridging has been used to explain the spread of information in a chain of members including three testable concepts: first, linearity assumes that individual “agent bees” that participate in the wave will be affected preferentially from the side of wave origin. The directed-trigger hypothesis addresses the coincidence of the individual property of trigger direction with the collective property of wave direction. Second, continuity describes the transfer of information without being stopped, delayed or re-routed. The active-neighbours hypothesis assumes coincidence between the direction of the majority of shimmering-active neighbours and the trigger direction of the agents. Third, the graduality hypothesis refers to the interaction between an agent and her active neighbours, assuming a proportional relationship in the strength of abdomen flipping of the agent and her previously active neighbours. Shimmering waves provoked by dummy wasps were recorded with high-resolution video cameras. Individual bees were identified by 3D-image analysis, and their strength of abdominal flipping was assessed by pixel-based luminance changes in sequential frames. For each agent, the directedness of wave propagation was based on wave direction, trigger direction, and the direction of the majority of shimmering-active neighbours. The data supported the bucket bridging hypothesis, but only for a small proportion of agents: linearity was confirmed for 2.5%, continuity for 11.3% and graduality for 0.4% of surface bees (but in 2.6% of those agents with high wave-strength levels). The complimentary part of 90% of surface bees did not conform to bucket bridging. This fuzziness is discussed in terms of self-organisation and evolutionary adaptedness in Giant honeybee colonies to respond to rapidly changing threats such as predatory wasps scanning in front of the nest.
Highlights
Shimmering behaviour in Giant honeybees (Apis dorsata) [1,2] involves a display of social waves with antipredatory impact [3,4,5,6,7,8,9]. These waves originate at discrete areas on the surface of the nests, where generator bees have been identified [10] as leaders in the general responsiveness to external cues
It is reasonable to assume that the mechanoceptive domain [16] of shimmering is important for colony-intrinsic communication [16], because the waves affect most of the colony members in the surface layer, but in all layers of the bee curtain [7,14]; they are even supposed to influence the bee curtain on the non-shimmering, opposite side of the comb [17]
Categorization of Active Participation in Shimmering Individual agent bees were identified on the surface of a Giant honeybee nest by stereoscopic imaging (Fig. 3A,B; [11]) for 66 manually selected shimmering waves (Fig. 3C)
Summary
Shimmering behaviour in Giant honeybees (Apis dorsata) [1,2] involves a display of social waves with antipredatory impact [3,4,5,6,7,8,9] These waves originate at discrete areas on the surface of the nests, where generator bees have been identified [10] as leaders in the general responsiveness to external cues. These generator bees raise their abdomen first and affect their nest mates around them to follow them in sequential order. It is reasonable to assume that the mechanoceptive domain [16] of shimmering is important for colony-intrinsic communication [16], because the waves affect most of the colony members in the surface layer, but in all layers of the bee curtain [7,14]; they are even supposed to influence the bee curtain on the non-shimmering, opposite side of the comb [17]
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