Abstract: The social hierarchies observed in natural systems often show a high degree of transitivity. Transitive hierarchies do not only require rank differentiation within pairs of individuals but also a higher level ordering of relations within the group. Several authors have suggested that the formation of linear hierarchies at the group level is an emergent property of individual behavioural rules, referred to as winner and loser effects. Winner and loser effects occur if winners of previous conflicts are more likely to escalate the current conflict, whereas the losers of previous conflicts are less likely to do so. According to this idea, an individual's position in a hierarchy may not necessarily reflect its fighting ability, but may rather result from arbitrary historical asymmetries, in particular the history of victories and defeats. However, if this is the case, it is difficult to explain from an evolutionary perspective why a low ranking individual should accept its subordinate status. Here we present a game theoretical model to investigate whether winner and loser effects giving rise to transitive hierarchies can evolve and under which conditions they are evolutionarily stable. The main version of the model focuses on an extreme case in which there are no intrinsic differences in fighting ability between individuals. The only asymmetries that may arise between individuals are generated by the outcome of previous conflicts. We show that, at evolutionary equilibrium, these asymmetries can be utilized for conventional conflict resolution. Several evolutionarily stable strategies are based on winner and loser effects and these strategies give rise to transitive hierarchies.

Abstract: We present the results of a series of computer simulations that examined the impact of winner, loser, and bystander effects on hierarchy formation in fused groups. These effects and their implications for hierarchy structure and aggressive interactions were first examined in small four-member groups. Subsequent to this, the two small groups were fused into a single larger group. Further interactions took place in this fused group, generating a new hierarchy. Our models demonstrate clearly that winner, loser, and bystander effects strongly influence both the structure and types of interactions that emerge from the fusion of smaller groups. Four conditions produced results in which the same general patterns were uncovered in pre- and postfusion groups: (1) winner effects alone, (2) bystander loser effects alone, (3) winner and bystander winner effects operating simultaneously, and (4) all four effects in play simultaneously. Outside this parameter space, hierarchy structure and the nature of aggressive interactions differed in pre- and postfusion groups. When only loser effects were in play, one of the two clear alphas from the prefused groups dropped in rank in the eight-member fused group. When bystander winner effects were in play, it was difficult to rank any of the eight individuals in the fused group, and players interacted almost exclusively with those that were not in their original four-member group. When loser and bystander loser effects operated simultaneously, two top-ranking individuals emerged in the fused groups, but the relative rank of the other players was difficult to assign.

Abstract: >We examined the “decision-making” process of aggressive interactions within a community of wild chimpanzees ( Pan troglodytes verus) in the Taï National Park, Côte d’Ivoire (West Africa). Costs and benefits were investigated for 876 dyadic aggressive interactions among 18 adults (including 4 independent adolescents) of either sex. An extended version of the Relational Model was developed to describe the dynamics of the “decision-making” process in Taï chimpanzees, which suggests that the net benefit determines the occurrence of conflicts. Both sexes fought more frequently for the resources that were most important to them, food for females and social contexts for males. Individuals used two different strategies according to their likelihood of winning the aggressive interaction, determined by the dominance relationship of the conflict partners. Dominant initiators had longer and more intense aggressive interactions, but they limited their social disadvantages by fighting non-cooperative partners. Subordinate initiators had shorter and less intense aggressive interactions, but risked more social costs, which they could reduce afterwards by reconciliation. Both strategies included a positive overall net benefit. The extended Relational Model fits the complexity of wild chimpanzee conflicts and allows for more flexibility in the “decision-making” compared to the original version.

Abstract: African wild dog (Lycaon pictus) pup vocalizations were studied in Hwange National Park, Zimbabwe for weeks 3 through 7 of the socialization period. Here we present the vocal repertoire, including the use of repetitive and mixed sounds, and investigate the extent to which the emerging sound system of Lycaon conforms to predicted design features of Morton's (MS) motivation-structural rules. Features of the pup sound system are highlighted by comparison with adults and other social canids.TAGSTARTBRTAGEND Data were collected at three den sites (litter sizes: 8, 8, and 9) of two study packs. A total of 1903 vocalizations were classified, and eight vocal classes and seven subclasses were identified. Although all sounds identified persist into adulthood, observations indicate a delayed onset in some vocal classes, including both the lowest (i.e. rumbles) and highest (i.e. twitters) frequency sounds. As predicted by the (MS) model, pups invested heavily in high frequency, harmonic care/social soliciting sounds (91%, N = 1586 unmixed vocalizations), however, no clear association between acoustic structure and sound repetition was found. Significantly more repetition was heard in all vocal classes with the exception of moans and barks. Intra-pack aggression is generally muted in this obligate social carnivore suggesting that repetition may be a low cost strategy to induce social outcomes and obtain food. The patterning of mixed vocalizations (N = 317) was consistent with the (MS) model. Given the high degree of cooperation necessary for individual survival, the predominant use of cross-mixed sounds may serve to minimize conflict as pups begin to form relationships with littermates and adults. Noisy/noisy sounds were exceptionally rare. Comparative data suggest a relationship between the early patterning of mixed sounds and species-specific social organization in canids.

Abstract: 1. The social relationships in a group of Icelandic horses without a mature stallion were studied. The horses were all familiar to each other. Mutual grooming and play relationships, spatial associations, dominance-subordinate relations and the effect of kinship on these relationships were analysed.TAGSTARTBRTAGEND 2. The social structure was clearly dominated by the behaviour of the adult mares. The horses preferred to form bonds within their social class (sex/age) and they kept close proximity with their friends. The group was effectively divided into two social subgroups, adult mares as one group and adult geldings and sub-adults as another group. The sub-adults and adult geldings formed associations, which were based on mutual grooming and play, while the adult mares did not play. Differences between the sexes were evident. Males played more than the females, had more playing partners and were more popular as playmates.TAGSTARTBRTAGEND 3. Aggression rates were low. The dominance hierarchy was linear. Adult mares ranked higher than adult geldings, sub-adults and the foals. Rank was significantly correlated with age. The closer the adult mares were in rank, the more they groomed with each other. Such relationships were not found amongst the other social group.TAGSTARTBRTAGEND 4. Kinship was calculated between all pairs of animals for up to 4 or 5 generations. Allogrooming and play frequencies and proximity were all positively correlated with kinship. Adult mares, which were close in the dominance hierarchy, were on average more related than those further apart.TAGSTARTBRTAGEND 5. The social relationships in the Icelandic herd were, to some extent, different from relationships reported from unmanaged and feral horse-herds with mature stallions and bachelors. Our results suggest that adult mares groom more in groups without a stallion. Furthermore, they have more preferred partners than in natural harems and their partners are other adult mares, not their weaned offspring as seems to be the case in feral herds. The sub-adults also seem to be more socially active in the absence of stallions. Interestingly, in the Icelandic group, the adult mares showed stallion like behaviours, like mounting and protecting foals. Only by studying the behaviour and the nature of the relationships of horses in groups of different compositions, can we expect to gain a comprehensive understanding about individual social strategies and cognitive capabilities of the species. Such knowledge is valuable for management and welfare of the horse.