Abstract: Indirect reciprocity1, 2, 3, 4, 5 is a key mechanism for the evolution of human cooperation. Our behaviour towards other people depends not only on what they have done to us but also on what they have done to others. Indirect reciprocity works through reputation5, 6, 7, 8, 9, 10, 11, 12, 13, 14, 15, 16, 17. The standard model of indirect reciprocity offers a binary choice: people can either cooperate or defect. Cooperation implies a cost for the donor and a benefit for the recipient. Defection has no cost and yields no benefit. Currently there is considerable interest in studying the effect of costly (or altruistic) punishment on human behaviour18, 19, 20, 21, 22, 23, 24, 25. Punishment implies a cost for the punished person. Costly punishment means that the punisher also pays a cost. It has been suggested that costly punishment between individuals can promote cooperation. Here we study the role of costly punishment in an explicit model of indirect reciprocity. We analyse all social norms, which depend on the action of the donor and the reputation of the recipient. We allow errors in assigning reputation and study gossip as a mechanism for establishing coherence. We characterize all strategies that allow the evolutionary stability of cooperation. Some of those strategies use costly punishment; others do not. We find that punishment strategies typically reduce the average payoff of the population. Consequently, there is only a small parameter region where costly punishment leads to an efficient equilibrium. In most cases the population does better by not using costly punishment. The efficient strategy for indirect reciprocity is to withhold help for defectors rather than punishing them.

Abstract: Group-living animals travel together to collectively exploit the resources of their environment. This study investigates how social relationships and individual temperament traits affect movement orders in domestic cattle and sheep. We analysed spontaneous group movements occurring at pasture after a resting period in a group of 15 18-month-old Charolais heifers and a group of 19 1-year-old Romane ewe-lambs. For each species, animals had similar social experience and no kinship ties. Before that, animals were observed within the group to establish their social status (e.g. dominance and preferential relationships, and sociability), then in individual tests in order to assess their emotional traits. In both species, most individuals could initiate a group movement but some individuals were more successful than others in recruiting the rest of the group. Ewe-lambs, and to a lesser extent heifers, held preferential positions during travel. We did not find any significant correlations in either species between animal order and their position in the dominance hierarchy (heifers: P = 0.438; ewe-lambs: P = 0.574) while individuals linked by preferential bonds frequently followed each other during group movements (heifers: P < 0.001; ewe-lambs: P < 0.001). With regard to social traits, heifers with a low cohesion index, and with a lower number of partners with whom they develop frequent affinitive interactions, acted more frequently as ‘first movers’ (P = 0.040 and 0.023, respectively), as well as did ewe-lambs with a high spatial independency index (P = 0.002). Ewe-lambs with the highest cohesion indices were more frequently observed in front of the group while moving halfway between departure and arrival (P = 0.028). We did not find significant correlations between individual positions during group movements and emotional traits such as reactivity, boldness and fearfulness. We conclude that preferential bonds and individual traits related to social dependence were more influential in spontaneous group movements at pasture than were emotional traits and dominance status.

Abstract: Mirror usage has been taken to indicate some degree of awareness in animals. Can pigs, Sus scrofa, obtain information from a mirror? When put in a pen with a mirror in it, young pigs made movements while apparently looking at their image. After 5 h spent with a mirror, the pigs were shown a familiar food bowl, visible in the mirror but hidden behind a solid barrier. Seven out of eight pigs found the food bowl in a mean of 23 s by going away from the mirror and around the barrier. Naïve pigs shown the same looked behind the mirror. The pigs were not locating the food bowl by odour, did not have a preference for the area where the food bowl was and did not go to that area when the food bowl was visible elsewhere. To use information from a mirror and find a food bowl, each pig must have observed features of its surroundings, remembered these and its own actions, deduced relationships among observed and remembered features and acted accordingly. This ability indicates assessment awareness in pigs. The results may have some effects on the design of housing conditions for pigs and may lead to better pig welfare.

Abstract: Abstract  The ability to select the greater numerosity over another in small sets seems to stem from the calculation of which set contains more, and has been taken as evidence of a primordial representation at the roots of the primate numerical system. We tested 56 horses (Equus caballus) in a paradigm previously used with human infants and nonhuman primates. Horses saw two quantities paired in contrasts—2 versus 1, 3 versus 2, 6 versus 4 and a control for volume, 2 versus 1 big—and had to make a choice by snout touching the container holding the numerosity selected. The horses spontaneously selected the greater of the two quantities when the numerosities were small. These results add to evidence showing spontaneous quantity assessment in a variety of species.

Abstract: While viewing faces, human adults often demonstrate a natural gaze bias towards the left visual field, that is, the right side of the viewee’s face is often inspected first and for longer periods. Using a preferential looking paradigm, we demonstrate that this bias is neither uniquely human nor limited to primates, and provide evidence to help elucidate its biological function within a broader social cognitive framework. We observed that 6-month-old infants showed a wider tendency for left gaze preference towards objects and faces of different species and orientation, while in adults the bias appears only towards upright human faces. Rhesus monkeys showed a left gaze bias towards upright human and monkey faces, but not towards inverted faces. Domestic dogs, however, only demonstrated a left gaze bias towards human faces, but not towards monkey or dog faces, nor to inanimate object images. Our findings suggest that face- and species-sensitive gaze asymmetry is more widespread in the animal kingdom than previously recognised, is not constrained by attentional or scanning bias, and could be shaped by experience to develop adaptive behavioural significance.