Insect visuomotor delay adjustments in group flight support swarm cohesion

Flying insects routinely demonstrate coordinated flight in crowded assemblies despite strict communication and processing constraints. This study experimentally records multiple flying insects tracking a moving visual stimulus. System identification techniques are used to robustly identify the tracking dynamics, including a visuomotor delay. The population delay distributions are quantified for solo and group behaviors. An interconnected visual swarm model incorporating heterogeneous delays is developed, and bifurcation analysis and swarm simulation are applied to assess swarm stability under the delays. The experiment recorded 450 insect trajectories and quantified visual tracking delay variation. Solitary tasks showed a 30ms average delay and standard deviation of 50ms, while group behaviors show a 15ms average and 8ms standard deviation. Analysis and simulation indicate that the delay adjustments during group flight support swarm formation and center stability, and are robust to measurement noise. These results quantify the role of visuomotor delay heterogeneity in flying insects and their role in supporting swarm cohesion through implicit communication.