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Leadership behavior is a widespread phenomenon in animal groups on the move such as insect swarms, bird ﬂocks, ﬁsh shoals and herds ( , , , , , ,  ). Mathematical modeling based on the analysis of observational data is often used to study leadership behavior ( , , ). The rules of interaction of individuals have been inferred from observation. (For example, a zonal rule showed that the attraction and repulsion of neighbors for mosquitoﬁsh depends on the distance between them .) However, these proposed rules have not been tested empirically, as the live individuals are hard to control .