Supplementary Material from On the emergence of gravitational-like forces in insect swarms

Okubo (Okubo 1986 Adv. Biophys. 22, 1–94. (doi:10.1016/0065-227X(86)90003-1)) was the first to propose that insect swarms are analogous to self-gravitating systems. In the intervening years, striking similarities between insect swarms and self-gravitating systems have been uncovered. Nonetheless, experimental observations of laboratory swarms provide no conclusive evidence of long-range forces acting between swarming insects. The insects appear somewhat paradoxically to be tightly bound to the swarm while at the same time weakly coupled inside it. Here, I show how resultant centrally attractive gravitational-like forces can emerge from the observed tendency of insects to continually switch between two distinct flight modes: one that consists of low-frequency manoeuvers and one that consists of higher-frequency nearly harmonic oscillations conducted in synchrony with another insect. The emergent dynamics are consistent with ‘adaptive’ gravity models of swarming and with variants of the stochastic models of Okubo and Reynolds for the trajectories of swarming insects: models that are in close accord with a plethora of observations of unperturbed and perturbed laboratory swarms. The results bring about a radical change of perspective as swarm properties can now be attributed to known biological behaviours rather than to elusive physical influences.