How perceived threat increases synchronization in collectively moving animal groups

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Abstract
Nature is rich with many different examples of the cohesive motion of animals. Previous attempts to model collective motion have primarily focused on group behaviours of identical individuals. In contrast, we put our emphasis on modelling the contributions of different individual-level characteristics within such groups by using stochastic asynchronous updating of individual positions and orientations. Our model predicts that higher updating frequency, which we relate to perceived threat, leads to more synchronized group movement, with speed and nearest-neighbour distributions becoming more uniform. Experiments with three-spined sticklebacks (Gasterosteus aculeatus) that were exposed to different threat levels provide strong empirical support for our predictions. Our results suggest that the behaviour of fish (at different states of agitation) can be explained by a single parameter in our model: the updating frequency. We postulate a mechanism for collective behavioural changes in different environment-induced contexts, and explain our findings with reference to confusion and oddity effects.