Collective motion, where large numbers of individuals move synchronously together, is achieved when individuals adopt interaction rules that determine how they respond to their neighbors' movements and positions. These rules determine how group-living animals move, make decisions, and transmit information between individuals. Nonetheless, few studies have explicitly determined these interaction rules in moving groups, and very little is known about the interaction rules of fish. Here, we identify three key rules for the social interactions of mosquitofish (Gambusia holbrooki): (i) Attraction forces are important in maintaining group cohesion, while we find only weak evidence that fish align with their neighbor's orientation; (ii) repulsion is mediated principally by changes in speed; (iii) although the positions and directions of all shoal members are highly correlated, individuals only respond to their single nearest neighbor. The last two of these rules are different from the classical models of collective animal motion, raising new questions about how fish and other animals self-organize on the move.collective animal behavior | fish shoals | group motion | self-propelled particles | self-organization C ollective motion of animal groups occurs when multiple individuals move synchronously, producing large scale "flocking" patterns (1-5). Numerous models have been developed to describe patterns of collective motion in terms of interactions between individuals (6-9). These simulation models usually assume that individuals move at a constant speed and their interactions are mediated through direction changes (1). Often these models use zonal rules, where individuals move away from neighbors at close distances and align and/or move toward neighbors at greater distances. Interactions can be with either all neighbors within some zone (7) or with a set of n nearest neighbors (10). These and other models have succeeded in reproducing qualitatively similar patterns to those seen in the collective motion of animal groups in nature.It remains unclear, however, whether the interaction rules implemented in models are the ones used by animals. Indeed, many collective motion patterns observed in nature can be simulated by models using very different interaction rules (1). We are only now beginning to accumulate evidence about which interaction rules are used. There has been recent identification of zones of repulsion and alignment in surf scoters (11). The structure of starling flocks is consistent with topological interactions between the birds (10). Homing pigeons appear to have hierarchical interactions such that birds with higher route-following fidelity act as leaders (12)(13)(14)(15). Partridge showed that lateral line and vision are both important in producing directional alignment in Gadid fish (16). Nonetheless, there remain a large number of open questions about the interactions of fish. For example, do fish adopt attraction and alignment within distinct zones as purported in most models? How many neighbors do fish ...

How different levels of biological organization interact to shape each other's function is a central question in biology. One particularly important topic in this context is how individuals' variation in behaviour shapes group-level characteristics. We investigated how fish that express different locomotory behaviour in an asocial context move collectively when in groups. First, we established that individual fish have characteristic, repeatable locomotion behaviours (i.e. median speeds, variance in speeds and median turning speeds) when tested on their own. When tested in groups of two, four or eight fish, we found individuals partly maintained their asocial median speed and median turning speed preferences, while their variance in speed preference was lost. The strength of this individuality decreased as group size increased, with individuals conforming to the speed of the group, while also decreasing the variability in their own speed. Further, individuals adopted movement characteristics that were dependent on what group size they were in. This study therefore shows the influence of social context on individual behaviour. If the results found here can be generalized across species and contexts, then although individuality is not entirely lost in groups, social conformity and group-size-dependent effects drive how individuals will adjust their behaviour in groups.

We examine changes in interaction rules, predictability, and vigilance of x-ray tetras due to external food and alarm cues.

Despite the frequency with which mixed-species groups are observed in nature, studies of collective behaviour typically focus on single-species groups. Here, we quantify and compare the patterns of interactions between three fish species, threespine sticklebacks (Gasterosteus aculeatus), ninespine sticklebacks (Pungitius pungitius) and roach (Rutilus rutilus) in both single- and mixed-species shoals in the laboratory. Pilot data confirmed that the three species form both single- and mixed-species shoals in the wild. In our laboratory study, we found that single-species groups were more polarized than mixed-species groups, while single-species groups of threespine sticklebacks and roach were more cohesive than mixed shoals of these species. Furthermore, while there was no difference between the inter-individual distances between threespine and ninespine sticklebacks within mixed-species groups, there was some evidence of segregation by species in mixed groups of threespine sticklebacks and roach. There were differences between treatments in mean pairwise transfer entropy, and in particular we identify species-differences in information use within the mixed-species groups, and, similarly, differences in responses to conspecifics and heterospecifics in mixed-species groups. We speculate that differences in the patterns of interactions between species in mixed-species groups may determine patterns of fission and fusion in such groups.