“…Social groups are intrinsically heterogeneous [20], with members that may differ in leadership status [21], social rank [14,[20][21][22][23], sex [13,20,21,24,25], size [20], personality [20,26,27] or age [20,21,24], among other factors. This heterogeneity is likely to lead to individual differences in the risks and rewards of fighting [13,20,25,[28][29][30]. For example, in African lions (Panthera leo), there are consistent differences between individuals in their willingness to engage in territory disputes, as some 'laggard' females consistently hang back and exploit the fighting effort of others [26,30,31].…”