Intra- versus intergroup variance in collective behavior.

Animal collective motion arises from the intricate interactions between the natural variability among individuals, and the homogenizing effect of the group, working to generate synchronization and maintain coherence. Here, these interactions were studied using marching locust nymphs under controlled laboratory settings. A novel experimental approach compared single animals, small groups, and virtual groups composed of randomly shuffled real members. We found that the locust groups developed unique, group-specific behavioral characteristics, reflected in large intergroup and small intragroup variance (compared with the shuffled groups). Behavioral features that differed between single animals and groups, but not between group types, were classified as essential for swarm formation. Comparison with Markov chain models showed that individual tendencies and the interaction network among animals dictate the group characteristics. Deciphering the bidirectional interactions between individual and group properties is essential for understanding the swarm phenomenon and predicting large-scale swarm behaviors.