Group-living organisms that collectively 1 migrate range from cells and bacteria to human crowds, 2 and include swarms of insects, schools of fish and flocks 3 of birds or ungulates. Unveiling the behavioural and 4cognitive mechanisms by which these groups coordinate 5 their movements is a challenging task. These mecha-6 nisms take place at the individual scale and they can 7 be described as a combination of pairwise interactions 8 between individuals and interactions between these in-9 dividuals and the physical obstacles in the environment. 10 Thanks to the development of novel tracking techniques 11 that provide large and accurate data sets, the main 12 characteristics of individual and collective behavioural 13 patterns can be quantified with an unprecedented level 14 of precision. However, in a large number of works, social 15 interactions are usually described by force map meth-16 ods that only have a limited capacity of explanation and 17 prediction, being rarely suitable for a direct implemen-18 tation in a concise and explicit mathematical model. 19 Here, we present a general method to extract the in-20 teractions between individuals that are involved in the 21 coordination of collective movements in groups of organ-22 isms. We then apply this method to characterize social 23 interactions in two species of shoaling fish, the rummy-24 nose tetra (Hemigrammus rhodostomus) and the ze-25 brafish (Danio rerio), which both present a burst-and-26 coast motion. The detailed quantitative description of 27 microscopic individual-level interactions thus provides 28 predictive models of the emergent dynamics observed 29 at the macroscopic group-level. This method can be 30 applied to a wide range of biological and social systems.31behavioural changes correspond to significant variations 88 of the individual's heading that occur exactly at the 89 onset of the acceleration phase (i.e., the bursts). These 90 discrete behavioural decisions are called "kicks" [12,16].
91Other quantities such as the intensity of the accelera-92 tion in the direction perpendicular to the direction of 93 motion, or simply the turning direction (right or left), 94 can be used to detect behavioural changes. The task 95 is thus to put in relation the heading variation of a fo-96 cal fish δφ i with its state variables, that is, to find a 97 function δφ i (d, v, ψ, φ). 98 In this article, we first show that force maps, that 99 are widely used to describe the effects of social interac-100 tions on the behaviour of individuals, have important 101 limitations when it comes to model the interactions be-102 tween individuals. These limitations result mostly from 103 the limited number of variables that force maps can 104 handle, the difficulty of identifying intermediate contri-105 butions to behavioural patterns, and the difficulty of 106 distinguishing the effects of state variables from consti-107 tutive parameters. We then describe in detail a method 108 to analyse behavioural data obtained from digitized in-109 dividual trajectories. The m...