Abstract: Meeting the psychological needs of chimpanzees (Pan troglodytes) can be a challenge given their aggressiveness on the one hand and the complexity of their social lives on the other. It is unclear how to balance the need to provide opportunities for species-appropriate behavior against potential risks of injury chimpanzees may inflict on each other. This study evaluates the suggestion that simpler social environments protect chimpanzees from wounding. Over a two-year period all visible injuries to 46 adult males, 64 adult females, and 25 immature chimpanzees were recorded at the Yerkes Regional Primate Research Center. Approximately half of the subjects were mother-reared, and the rest were nursery-reared. Housing included compounds containing about 20 chimpanzees, interconnected indoor-outdoor runs for groups of up to 12 individuals, and smaller indoor-outdoor runs for pairs and trios. Annual wounding rates were calculated for serious wounds (extensive injuries and all those requiring veterinary intervention) as well as for minor wounds. Compound-housed chimpanzees incurred the highest level of minor wounding, but serious wounding levels were not affected by housing condition. Even with a period of dominance instability and elevated levels of wounding in one compound, compound chimpanzees were not injured more than those in smaller social groups over the long term. Nursery-reared females in moderate-sized groups were wounded more than mother-reared females. Also, nursery-reared males and females were wounded less often when paired with mother-reared companions. Overall, this study indicates that maintaining chimpanzees in pairs and trios would not be an effective means for reducing injuries. The management of wounding in chimpanzee colonies is influenced more by the sex and rearing composition of a colony.

Abstract: Feral horses are social animals, which have to rely on survival strategies centered on the formation of cohesive social bonds within their bands. Many problems in the husbandry of social animals such as horses, are due to the fact that the limits of their adaptive abilities are exceeded. Evidence suggests that the fundamental social characteristics of domestic horses have remained relatively unchanged. The social structure, social strategies and social interactions were investigated (3 non-consecutive years, 24 hr per day for several weeks) in long term established groups of domestic horses (mares and geldings of all ages) and a few small introduced groups, kept in (semi)natural environments. The general aim was to investigate the social needs of domestic horses. The social life of domestic horses was characterised by long lasting bonds with preferred partners which were established and maintained by allogrooming, play, proximity and dominance behaviours. Bonding partners were mainly found within the same sex-age group, but adult geldings also bonded with sub-adult mares and geldings. Adult mares were clustered in a group, while the other animals formed a second group. Among the adult mares, subgroups according to reproductive state were formed. Individuals regulated their social network by interfering with interactions between other members of the herd, which in itself is complex. An intervention is a behavioural action of one animal that actively interferes with an ongoing interaction between a dyad with the apparent aim of altering that interaction. This was verified by post-hoc analyses of disturbed and undisturbed interactions. Interventions in allogrooming or play were performed significantly more often when at least one member of the initial dyad was a preferred partner of, or familiar to (within the small introduced bands) the intervener. The stronger the preferred association in allogrooming between the intervener and member(s) of the initial dyad, the higher the probability the intervener would displace one initial member and continue allogrooming with the other. Just five behaviours were extracted which reliably reflected the dominance relations among horses. Aggression with the hind quarters was used both offensively and defensively and therefore not suitable as a reliable parameter. Individual dominance relationships were related to social experience. The implications of these findings for horse husbandry were assessed. It is argued that the execution of affiliative behaviours may be rewarding in itself, and therefore always will be a highly motivated behaviour. It is shown that social positive physical interactions (allogrooming, play) with other horses is an ethological need and therefore indispensable in modern husbandry systems. Ethological needs are so important for the animal that husbandry systems that lack the possibilities to execute such behaviours will cause chronic stress. It is concluded that all horses need physical social contact, and that horses, which lack appropriate social learning experiences during ontogeny, may be hampered in their social functioning later in life. Solutions for problems, including dominance problems, in individual social housing and group housing are presented.

Abstract: tEye and limb preferences were scored in the closest undomesticated relative of Equuscaballus using the same methods as used previously to study laterality in feral horses.Observations were made of 33 Przewalski horses (Equus ferus przewalskii) (male N = 20,female N = 13) living under natural social conditions on a large reserve in France. Signifi-cant left-eye/side biases were found in agonistic interactions within harem bands (M ± SEbias to left 58% ± 0.01 for threats, P < 0.001; 68% ± 0.05 for attacks; P < 0.001) and in stallionfights (threats, 52% ± 0.01 left, P < 0.001; attacks, 63% ± 0.02 left, P < 0.001): as many as 80%of the horses were significantly lateralized in attack responses within harem bands. Lat-erality of vigilance was measured as lifting up the head from grazing and turning it to theleft or right side: a directional bias to the left was found (M ± SE 53% ± 0.02 left, P < 0.001).Side bias in reactivity was calculated as the percent of head lifts above the level of thewithers on the left or right side and this was also left side biased (M ± SE 73% ± 0.03 left,P < 0.001). These results indicate right-hemisphere specialization for control of aggressionand responses to novelty. The left bias in attack scores within harem bands was strongerin males than females (P = 0.024) and in immature than adult horses (P = 0.032). Immaturehorses were also more strongly lateralized than adults in vigilance scores (P = 0.022), whichmay suggest that experience reduces these side biases. Our results show that Przewalskihorses exhibit left eye preferences, as do feral horses, and do so even more strongly thanferal horses. Considering feral and Przewalski horses together, we deduce that ancestralhorses had similar lateral biases. Also similar to feral horses, the Przewalski horses showedno significant forelimb preference at the group level or in the majority of horses at theindividual level, confirming the hypothesis that previously reported limb preferences indomestic breeds are entrained or generated by breed-specific selection.

Abstract: Observational field studies were conducted on two remote populations of feral horses in Australia to determine whether lateralization is a characteristic of Equus caballus as a species or results from handling by humans. Group 1 had been feral for two to five generations and Group 2 for 10–20 generations. In both groups, left-side biases were present during agonistic interactions and in reactivity and vigilance. Therefore, as in other vertebrates, the right hemisphere appears to be specialized to control agonistic behaviour and responses to potential threats. The leftwards bias was stronger in measures of behaviour involving more aggression and reactivity. Preferences to place one forelimb in front of the other during grazing were also determined. No population bias of forelimb preference was found, suggesting that such limb preferences present in domestic horses may be entrained. Since stronger individual limb preferences were found in immature than in adult feral horses, limb preference may be modified by maturation or experience in the natural habitat. Stronger limb preference was associated significantly with elevated attention to the environment but only in younger feral horses. No sex differences in lateralization were found. The findings are evidence that horses show visual lateralization, as in other vertebrates, not dependent on handling by humans. Limb preference during grazing, by contrast, does appear to depend on experience.

Abstract: Remembering individual identities is part of our own everyday social life. Surprisingly, this ability has recently been shown in two social insects. While paper wasps recognize each other individually through their facial markings, the ant, Pachycondyla villosa, uses chemical cues. In both species, individual recognition is adaptive since it facilitates the maintenance of stable dominance hierarchies among individuals, and thus reduces the cost of conflict within these small societies. Here, we investigated individual recognition in Pachycondyla ants by quantifying the level of aggression between pairs of familiar or unfamiliar queens over time. We show that unrelated founding queens of P. villosa and Pachycondyla inversa store information on the individual identity of other queens and can retrieve it from memory after 24h of separation. Thus, we have documented for the first time that long-term memory of individual identity is present and functional in ants. This novel finding represents an advance in our understanding of the mechanism determining the evolution of cooperation among unrelated individuals.