Thermoregulatory morphodynamics of honeybee swarm clusters

During reproductive swarming, honeybee clusters of more than 10,000 individuals that hang from structures in the environment (e.g. tree branches) are exposed to diurnal variations in ambient temperature for up to a week. Swarm clusters collectively modulate their morphology in response to these variations (i.e. expanding/contracting in response to heating/cooling) to maintain their internal temperature within a tolerable range and to avoid exhausting their honey stores prematurely. To understand the spatiotemporal aspects of thermoregulatory morphing, we measured the change in size, shape and internal temperature profiles of swarm clusters in response to dynamic temperature ramp perturbations. Swarm clusters showed a two-fold variation in their volume/density when heated from 15°C to 30°C. However, they did not reach an equilibrium size or shape when held at 30°C for 5 h, long after the core temperature of the cluster had stabilized. Furthermore, the changes in cluster shape and size were hysteretic, contracting in response to cooling faster than expanding in response to heating. Although the base contact diameter of the cluster increased continuously when the swarm was heated, the change in length of the swarm (base to tip) over time was non-monotonic. Consequently, the aspect ratio of the swarm fluctuated continuously even when held at a constant temperature. Taken together, our results quantify the hysteretic and anisotropic morphological responses of swarm clusters to ambient temperature variations while suggesting that both mechanical constraints and heat transfer govern their thermoregulatory morphodynamics.