Hierarchical organization is widespread in the societies of humans and other animals, both in social structure and in decision-making contexts. In the case of collective motion, the majority of case studies report that dominant individuals lead group movements, in agreement with the common conflation of the terms "dominance" and "leadership." From a theoretical perspective, if social relationships influence interactions during collective motion, then social structure could also affect leadership in large, swarm-like groups, such as fish shoals and bird flocks. Here we use computer-vision-based methods and miniature GPS tracking to study, respectively, social dominance and in-flight leader-follower relations in pigeons. In both types of behavior we find hierarchically structured networks of directed interactions. However, instead of being conflated, dominance and leadership hierarchies are completely independent of each other. Although dominance is an important aspect of variation among pigeons, correlated with aggression and access to food, our results imply that the stable leadership hierarchies in the air must be based on a different set of individual competences. In addition to confirming the existence of independent and context-specific hierarchies in pigeons, we succeed in setting out a robust, scalable method for the automated analysis of dominance relationships, and thus of social structure, applicable to many species. Our results, as well as our methods, will help to incorporate the broader context of animal social organization into the study of collective behavior.collective animal behavior | hierarchy | high-throughput ethology | leadership | dominance network F or a group of humans or nonhuman animals, networks can be constructed from a number of different types of interaction and across a range of contexts, including association, aggression, courtship, and leadership (1-4). This aspect of interaction networks raises the question as to whether network structure is maintained across contexts because of stable relationships or underlying individual differences, or whether network structure reorganizes in every new situation, where the same individuals may have different competences. For example, does social dominance routinely endow individuals with leadership roles within the group? By leadership, we refer to an individual's degree of influence over a group's decision: in the case of collective travel, this largely concerns the timing or direction of the group's movements (5). Dominance signifies the consistent winning of agonistic interactions (6). Most studies of the relationship between leadership and social structure have focused on mammals, especially primates. In species with highly asymmetrical dominance relationships, such as gray wolves (7), mountain gorillas (8), and chacma baboons (9), there is a consistent trend for dominants to lead. This effect may be mediated by factors other than dominance, such as the central position of dominants in the association network (10) or their greater metabo...

Travelling in groups gives animals opportunities to share route information by following cues from each other's movement. The outcome of group navigation will depend on how individuals respond to each other within a flock, school, swarm or herd. Despite the abundance of modelling studies, only recently have researchers developed techniques to determine the interaction rules among real animals. Here, we use high-resolution GPS (global positioning system) tracking to study these interactions in pairs of pigeons flying home from a familiar site. Momentary changes in velocity indicate alignment with the neighbour's direction, as well as attraction or avoidance depending on distance. Responses were stronger when the neighbour was in front. From the flocking behaviour, we develop a model to predict features of group navigation. Specifically, we show that the interactions between pigeons stabilize a side-by-side configuration, promoting bidirectional information transfer and reducing the risk of separation. However, if one bird gets in front it will lead directional choices. Our model further predicts, and observations confirm, that a faster bird (as measured from solo flights) will fly slightly in front and thus dominate the choice of homing route. Our results explain how group decisions emerge from individual differences in homing flight behaviour.

A key question in collective behavior is how individual differences structure animal groups, affect the flow of information, and give some group members greater weight in decisions. Depending on what factors contribute to leadership, despotic decisions could either improve decision accuracy or interfere with swarm intelligence. The mechanisms behind leadership are therefore important for understanding its functional significance. In this study, we compared pigeons' relative influence over flock direction to their solo flight characteristics. A pigeon's degree of leadership was predicted by its ground speeds from earlier solo flights, but not by the straightness of its previous solo route. By testing the birds individually after a series of flock flights, we found that leaders had learned straighter homing routes than followers, as we would expect if followers attended less to the landscape and more to conspecifics. We repeated the experiment from three homing sites using multiple independent flocks and found individual consistency in leadership and speed. Our results suggest that the leadership hierarchies observed in previous studies could arise from differences in the birds' typical speeds. Rather than reflecting social preferences that optimize group decisions, leadership may be an inevitable consequence of heterogeneous flight characteristics within self-organized flocks. We also found that leaders learn faster and become better navigators, even if leadership is not initially due to navigational ability. The roles that individuals fall into during collective motion might therefore have far-reaching effects on how they learn about the environment and use social information.

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For animals that travel in groups, the directional choices of conspecifics are potentially a rich source of information for spatial learning. In this study, we investigate how the opportunity to follow a locally experienced demonstrator affects route learning by pigeons over repeated homing flights. This test of social influences on navigation takes advantage of the individually distinctive routes that pigeons establish when trained alone. We found that pigeons learn routes just as effectively while flying with a partner as control pigeons do while flying alone. However, rather than learning the exact route of the demonstrator, the paired routes shifted over repeated flights, which suggests that the birds with less local experience also took an active role in the navigational task. The efficiency of the original routes was a key factor in how far they shifted, with less efficient routes undergoing the greatest changes. In this context, inefficient routes are unlikely to be maintained through repeated rounds of social transmission, and instead more efficient routes are achieved because of the interaction between social learning and information pooling.