Abstract: Summary For animals living in mixed-sex social groups, females who form strong social bonds with other females live longer and have higher offspring survival [1–3]. These bonds are highly nepotistic, but sometimes strong bonds may also occur between unrelated females if kin are rare [2, 3] and even among postdispersal unrelated females in chimpanzees and horses [4, 5]. Because of fundamental differences between the resources that limit reproductive success in females (food and safety) and males (fertilizations), it has been predicted that bonding among males should be rare and found only for kin and among philopatric males [6] like chimpanzees [7–9]. We studied social bonds among dispersing male Assamese macaques (Macaca assamensis) to see whether males in multimale groups form differentiated social bonds and whether and how males derive fitness benefits from close bonds. We found that strong bonds were linked to coalition formation, which in turn predicted future social dominance, which influenced paternity success. The strength of males' social bonds was directly linked to the number of offspring they sired. Our results show that differentiated social relationships exert an important influence on the breeding success of both sexes that transcends contrasts in relatedness.

Abstract: The Y chromosome directly reflects male genealogies, but the extremely low Y chromosome sequence diversity in horses has prevented the reconstruction of stallion genealogies [1, 2]. Here, we resolve the first Y chromosome genealogy of modern horses by screening 1.46 Mb of the male-specific region of the Y chromosome (MSY) in 52 horses from 21 breeds. Based on highly accurate pedigree data, we estimated the de novo mutation rate of the horse MSY and showed that various modern horse Y chromosome lineages split much later than the domestication of the species. Apart from few private northern European haplotypes, all modern horse breeds clustered together in a roughly 700-year-old haplogroup that was transmitted to Europe by the import of Oriental stallions. The Oriental horse group consisted of two major subclades: the Original Arabian lineage and the Turkoman horse lineage. We show that the English Thoroughbred MSY was derived from the Turkoman lineage and that English Thoroughbred sires are largely responsible for the predominance of this haplotype in modern horses.

Abstract: Summary For humans, facial expressions are important social signals, and how we perceive specific individuals may be influenced by subtle emotional cues that they have given us in past encounters. A wide range of animal species are also capable of discriminating the emotions of others through facial expressions [1, 2, 3, 4, 5], and it is clear that remembering emotional experiences with specific individuals could have clear benefits for social bonding and aggression avoidance when these individuals are encountered again. Although there is evidence that non-human animals are capable of remembering the identity of individuals who have directly harmed them [6, 7], it is not known whether animals can form lasting memories of specific individuals simply by observing subtle emotional expressions that they exhibit on their faces. Here we conducted controlled experiments in which domestic horses were presented with a photograph of an angry or happy human face and several hours later saw the person who had given the expression in a neutral state. Short-term exposure to the facial expression was enough to generate clear differences in subsequent responses to that individual (but not to a different mismatched person), consistent with the past angry expression having been perceived negatively and the happy expression positively. Both humans were blind to the photograph that the horses had seen. Our results provide clear evidence that some non-human animals can effectively eavesdrop on the emotional state cues that humans reveal on a moment-to-moment basis, using their memory of these to guide future interactions with particular individuals.

Abstract: Summary The transmission of cultural knowledge requires learners to identify what relevant information to retain and selectively imitate when observing others' skills. Young human infants--without relying on language or theory of mind--already show evidence of this ability. If, for example, in a communicative context, a model demonstrates a head action instead of a more efficient hand action, infants imitate the head action only if the demonstrator had no good reason to do so, suggesting that their imitation is a selective, interpretative process [1]. Early sensitivity to ostensive-communicative cues and to the efficiency of goal-directed actions is thought to be a crucial prerequisite for such relevance-guided selective imitation [2]. Although this competence is thought to be human specific [2], here we show an analog capacity in the dog. In our experiment, subjects watched a demonstrator dog pulling a rod with the paw instead of the preferred mouth action. In the first group, using the “inefficient” action was justified by the model's carrying of a ball in her mouth, whereas in the second group, no constraints could explain the demonstrator's choice. In the first trial after observation, dogs imitated the nonpreferred action only in the second group. Consequently, dogs, like children, demonstrated inferential selective imitation.

Abstract: Summary Although animals (particularly tool-users) are capable of solving physical tasks in the laboratory and the degree to which they understand them in terms of their underlying physical forces is a matter of contention. Here, using a new paradigm, the two-trap tube task, we report the performance of non-tool-using rooks. In contrast to the low success rates of previous studies using trap-tube problems , , and , seven out of eight rooks solved the initial task, and did so rapidly. Instead of the usual, conceptually flawed control, we used a series of novel transfer tasks to test for understanding. All seven transferred their solution across a change in stimuli. However, six out of seven were unable to transfer to two further tasks, which did not share any one visual constant. One female was able to solve these further transfer tasks. Her result is suggestive evidence that rooks are capable of sophisticated physical cognition, if not through an understanding of unobservable forces and , perhaps through rule abstraction. Our results highlight the need to investigate cognitive mechanisms other than causal understanding in studying animal physical cognition.