Abstract: There are numerous reports of novel learned behaviour patterns in animal populations, yet the factors influencing the invention and spread of these innovations remain poorly understood. Here we investigated to what extent the pattern of spread of innovations in captive groups of starlings, Sturnus vulgaris, could be predicted by knowledge of individual and social group variables, including association patterns, social rank orders, measures of neophobia and asocial learning performance. We presented small groups of starlings with a series of novel extractive foraging tasks and recorded the latency for each bird to contact and solve each task, as well as the orders of contacting and solving. We then explored which variables best predicted the observed diffusion patterns. Object neophobia and social rank measures characterized who was the first of the group to contact the novel foraging tasks, and the subsequent spread of contacting tasks was associated with latency to feed in a novel environment. Asocial learning performance, measured in isolation, predicted who was the first solver of the novel foraging tasks in each group. Association patterns did not predict the spread of solving. Contact latency and solving duration were negatively correlated, consistent with social learning underlying the spread of solving. Our findings indicate that we can improve our understanding of the diffusion dynamics of innovations in animal groups by investigating group-dependent and individual variables in combination. We introduce novel methods for exploring predictors of the origin and spread of behavioural innovations that could be widely applied.

Abstract: The social intelligence hypothesis argues that competition and cooperation among individuals have shaped the evolution of cognition in animals. What do we mean by social cognition? Here we suggest that the building blocks of social cognition are a suite of skills, ordered roughly according to the cognitive demands they place upon individuals. These skills allow an animal to recognize others by various means; to recognize and remember other animals' relationships; and, perhaps, to attribute mental states to them. Some skills are elementary and virtually ubiquitous in the animal kingdom; others are more limited in their taxonomic distribution. We treat these skills as the targets of selection, and assume that more complex levels of social cognition evolve only when simpler methods are inadequate. As a result, more complex levels of social cognition indicate greater selective pressures in the past. The presence of each skill can be tested directly through field observations and experiments. In addition, the same methods that have been used to compare social cognition across species can also be used to measure individual differences within species and to test the hypothesis that individual differences in social cognition are linked to differences in reproductive success.

Abstract: Abstract There is increasing interest in keeping horses in groups, but progress is hampered by a lack of knowledge about which horses can and should be kept together. Therefore, our objective was to investigate the effect of group composition on the occurrence of injuries among horses, the ease of removing horses from groups and horses’ reactivity to a fearful stimulus. Using a matched case control design, 61 groups of horses were studied in Denmark, Norway, Finland and Sweden. They were allocated into groups of similar or different age and sex or where membership changed regularly or remained stable. Injuries were recorded before mixing the horses into treatment groups, the day after mixing and four weeks later. Reactivity of horses to a moving novel object and the behaviour of a horse being removed from its group and the reactions of other group members towards this horse and the handler were evaluated. It was hypothesized that a more socially variable group composition has beneficial effects on behaviour, ease of handling and reducing reactivity whereas frequent changes in group composition has negative consequences, resulting in more injuries. We found that differences in treatment effects were mainly related to breed, rather than group composition. Icelandic horses reacted less to the movement of the novel object (P = 0.007) and approached it more afterwards (P = 0.04). They also had fewer new injuries than warmbloods following mixing (P < 0.001) and fewer than all other groups 4 weeks later (P < 0.01). Most new injuries after mixing were minor and recorded on the horse’s head, chest, hind legs and rump. In conclusion, variations in sex and age composition of the group had little effect on injury level, reactivity and ease of handling compared to the general effect of breed. Concerns about the risk of severe injuries associated with keeping horses in groups are probably overestimated. Thus, we propose that horses can be successfully kept in groups of different sex and age composition.

Abstract: We recently observed the spreading of a novel tradition in a flock of semiferal greylag geese, Anser anser: an increasing number of individuals began to bite and chew the stems of butterbur, Petasites hybridus. Because this behaviour spread particularly fast within families, social learning seemed to be involved. We therefore designed an experiment with hand-reared goslings, which were socially imprinted on humans, to investigate whether and how the observation of an experienced tutor affects the acquisition of a novel skill. Goslings had to open the gliding lid of a box to get at a food reward. To each of seven hand-reared observers a human tutor demonstrated where and how to open the lid, whereas seven controls remained untutored. All observers learned to perform the task but only one of the controls succeeded. The observers explored more often at the position shown by the tutor than elsewhere and seemingly learned by trial and error. In contrast, control birds explored primarily at positions that did not allow them to open the box. These results indicate that in greylag goslings the observation of an experienced model facilitates the learning of an operant task. We conclude that stimulus enhancement followed by operant conditioning were the mechanisms involved, which may have accounted for the fast spread of the stem-chewing tradition between family members.

Abstract: Aggression can generate anxiety, create uncertainty about its aftermath and jeopardise social relationships. Postconflict interactions serve as conflict management strategies to mitigate these consequences. Whereas postconflict interactions are well characterized in many animals, their functions are still insufficiently investigated. Four functional hypotheses have been proposed: stress reduction, relationship repair, self-protection and benign intent. We aimed to test these hypotheses in females of a tolerant macaque species, the crested macaque, Macaca nigra, under natural conditions, for three postconflict interactions: reconciliation, affiliation and aggression with third parties. Our results provide meaningful contrasts compared with findings in other species. We found no evidence that aggression had consequences for individuals' behavioural indicators of anxiety, although it increased the likelihood of secondary aggression with third parties. There was little evidence for the stress reduction hypothesis as the occurrence of any of the three postconflict interactions investigated had little effect on the measured behavioural indicators of anxiety. Conflict and dyad characteristics also had limited influence on anxiety. The relationship repair function was only partly validated: dyads with stronger bonds or that exchanged more support did not reconcile more often, but dyads with attributes related to the symmetry, stability and predictability (i.e. security) within relationships did. Patterns of initiation and directionality of postconflict interactions in this study population suggest that reconciliation may constitute the signalling of appeasement and benign intent. Furthermore, we found that aggression towards third parties may serve as a source of self-protection and reassertion of the females' social status. The distinctive pattern of postconflict management strategies revealed in wild female crested macaques appears to be related to their typically tolerant social style. These results demonstrate the usefulness of concomitantly studying aggression, postconflict interactions and their functions, to understand conflict management strategies comprehensively, while taking into account the level of social tolerance characterizing the studied society.