Abstract: In many animal groups, certain individuals consistently appear at the forefront of coordinated movements [1-4]. How such leaders emerge is poorly understood [5, 6]. Here, we show that in pairs of sticklebacks, Gasterosteus aculeatus, leadership arises from individual differences in the way that fish respond to their partner's movements. Having first established that individuals differed in their propensity to leave cover in order to look for food, we randomly paired fish of varying boldness, and we used a Markov Chain model to infer the individual rules underlying their joint behavior. Both fish in a pair responded to each other's movements-each was more likely to leave cover if the other was already out and to return if the other had already returned. However, we found that bolder individuals displayed greater initiative and were less responsive to their partners, whereas shyer individuals displayed less initiative but followed their partners more faithfully; they also, as followers, elicited greater leadership tendencies in their bold partners. We conclude that leadership in this case is reinforced by positive social feedback.

Abstract: Leadership is not an inherent quality of animal groups that show directional locomotion. However, there are other factors that may be responsible for the occurrence of leadership in fish shoals, such as individual differences in nutritional state between group members. It appears that front fish have a strong influence on directional shoal movements and that individuals that occupy such positions are often characterised by larger body lengths and lower nutritional state. Potential interactions between the two factors and their importance for positioning within shoals need further attention. Initiation of directional movement in stationary shoals and position preferences in mobile shoals need to be addressed separately because they are potentially subject to different constraints. Individuals that initiate a swimming direction may not necessarily be capable of the sustained high swimming performance required to keep the front position or have the motivation to do so, for that matter. More empirical and theoretical work is necessary to look at the factors controlling positioning behaviour within shoals, as well as overall shoal shape and structure. Tracking of marked individuals whose positioning behaviour is monitored over extended time periods of hours or days would be useful. There is an indication that shoal positions are rotated by individuals according to their nutritional needs, with hungry fish occupying front positions only for as long as necessary to regain their nutritional balance. This suggests that shoal members effectively take turns at being leaders. There is a need for three-dimensional recordings of shoaling behaviour using high-speed video systems that allow a detailed analysis of information transfer in shoals of different size. The relationship between leadership and shoal size might provide an interesting field for future research. Most studies to date have been restricted to shoals of small and medium size and more information on larger shoals would be useful.

Abstract: The term `leadership' has been used in several different senses, resulting in very different ways of identifying leaders and apparently inconsistent conclusions on how leadership is determined in herbivores. We therefore propose the following definitions: (i) a leader is the individual that is consistently the one who initiates long-distance, spontaneous group movements toward a new feeding site and (ii) long-distance spontaneous group movements are movements which happen when an animal changes activity and location and is immediately followed by a similar change in activity and location by other members of the group. Using these definitions, we tested for consistency of movement order across time and situation within a group of fifteen 2-year-old heifers. We found that the same individual was recorded as the very first animal in 48% of movements toward a new feeding site and could therefore be identified as the `leader'. We also showed that movement order when the animals entered an experimental plot, or progressed slowly through the field during a grazing bout, did not produce the same result. This method, which enables us to identify leaders in groups of animals at pasture, should improve our knowledge of how leadership is determined in grazing herbivores.

Abstract: This paper reviews the literature on leadership in vertebrate groups, including recent work on human groups, before presenting the results of three new experiments looking at leadership and decision making in small and large human groups. In experiment 1, we find that both group size and the presence of uninformed individuals can affect the speed with which small human groups (eight people) decide between two opposing directional preferences and the likelihood of the group splitting. In experiment 2, we show that the spatial positioning of informed individuals within small human groups (10 people) can affect the speed and accuracy of group motion. We find that having a mixture of leaders positioned in the centre and on the edge of a group increases the speed and accuracy with which the group reaches their target. In experiment 3, we use large human crowds (100 and 200 people) to demonstrate that the trends observed from earlier work using small human groups can be applied to larger crowds. We find that only a small minority of informed individuals is needed to guide a large uninformed group. These studies build upon important theoretical and empirical work on leadership and decision making in animal groups.

Abstract: Understanding how groups of individuals with different motives come to daily decisions about the exploitation of their environment is a key question in animal behaviour. While interindividual differences are often seen only as a threat to group cohesion, growing evidence shows that they may to some extent facilitate effective collective action. Recent studies suggest that personality differences influence how individuals are attracted to conspecifics and affect their behaviour as an initiator or a follower. However, most of the existing studies are limited to a few taxa, mainly social fish and arthropods. Horses are social herbivores that live in long-lasting groups and show identifiable personality differences between individuals. We studied a group of 38 individuals living in a 30-ha hilly pasture. Over 200 h, we sought to identify how far individual differences such as personality and affinity distribution affect the dynamic of their collective movements. First, we report that individuals distribute their relationships according to similar personality and hierarchical rank. This is the first study that demonstrates a positive assortment between unrelated individuals according to personality in a mammal species. Second, we measured individual propensity to initiate and found that bold individuals initiated more often than shy individuals. However, their success in terms of number of followers and joining duration did not depend on their individual characteristics. Moreover, joining process is influenced by social network, with preferred partners following each other and bolder individuals being located more often at the front of the movement. Our results illustrate the importance of taking into account interindividual behavioural differences in studies of social behaviours.