Abstract: Most animals must learn some of the behaviours in their repertoire, and some must learn most. Although learning is often thought of as an individual exercise, in nature much learning is social, i.e. under the influence of conspecifics. Social learners acquire novel information or skills faster and at lower cost, but risk learning false information or useless skills. Social learning can be divided into learning from social information and learning through social interaction. Different species have different mechanisms of learning from social information, ranging from selective attention to the environment due to the presence of others to copying of complete motor sequences. In vertical (or oblique) social learning, naïve individuals often learn skills or knowledge from parents (or other adults), whereas horizontal social learning is from peers, either immatures or adults, and more often concerns eavesdropping and public information use. Because vertical social learning is often adaptive, maturing individuals often have a preference for it over individual exploration. The more cognitively demanding social learning abilities probably evolved in this context, in lineages where offspring show long association with parents and niches are complex. Because horizontal learning can be maladaptive, especially when perishable information has become outdated, animals must decide when to deploy social learning. Social learning of novel skills can lead to distinct traditions or cultures when the innovations are sufficiently rare and effectively transmitted socially. Animal cultures may be common but to date taxonomic coverage is insufficient to know how common. Cultural evolution is potentially powerful, but largely confined to humans, for reasons currently unknown. A general theory of culture is therefore badly needed.

Abstract: Breeding groups with multiple stallions occur sympatrically with single-stallion breeding groups in feral horse, Equus caballus, populations. Mutualism and reciprocal altruism between stallions have been proposed to explain the origin and functioning of multistallion bands. However, empirical support for these hypotheses is contradictory and incomplete. Furthermore, there are no explicit tests of the predictions that each hypothesis makes about stallion behaviour and social structure. We compared nine multistallion and 18 single-stallion bands in the Kaimanawa Ranges, New Zealand. Compared with agonistic behaviours, affiliative behaviours were relatively unimportant in the relationships between stallions within bands. The number of stallions in the band did not have a positive influence on mare group size, stability, home range quality or reproductive success in bands. Furthermore, there was a positive relationship between aggression ('intolerance') by the dominant towards subordinate stallions and the subordinates' effort in mare group defence ('helping') but a negative relationship between helping effort by subordinates and their proximity to, and mating with, the bands' mares. Therefore, the predictions of the mutualism and reciprocal altruism hypotheses were not supported. Indeed, for some of the predictions we found the opposite outcomes to be true. Multistallion bands had significantly poorer reproductive success, and dominant stallions were less tolerant of subordinates that helped most and reduced their access to mares. Nevertheless, in all other respects Kaimanawa stallions in multistallion bands behaved like those described elsewhere. Thus, we reject cooperative hypotheses for multimale breeding groups in horses and discuss the mate parasitism and consort hypotheses as better alternatives. Copyright 2000 The Association for the Study of Animal Behaviour.

Abstract: We used a cooperative pulling task to examine proximate aspects of cooperation in captive brown capuchin monkeys, Cebus apella. Specifically, our goal was to determine whether capuchins can learn the contingency between their partner's participation in a task and its successful completion. We examined whether the monkeys visually monitored their partners and adjusted pulling behaviour according to their partner's presence. Results on five same-sex pairs of adults indicate that (1) elimination of visual contact between partners significantly decreased success, (2) subjects glanced at their partners significantly more in cooperative tests than in control tests in which no partner-assistance was needed, and (3) they pulled at significantly higher rates when their partner was present rather than absent. Therefore, in contrast to a previous report by Chalmeau et al. (1997, Animal Behaviour, 54, 1215-1225), cooperating capuchins do seem able to take the role of their partner into account. However, the type of task used may be an important factor affecting the level of coordination achieved. Copyright 2000 The Association for the Study of Animal Behaviour.

Abstract: When foraging together, animals are often observed to feed from food discoveries of others. The producer-scrounger (PS) game predicts how frequently this phenomenon of food parasitism should occur. The game assumes: (1) at any moment all individuals can unambiguously be categorized as either playing producer (searching for undiscovered food resources) or scrounger (searching for exploitation opportunities), and (2) the payoffs received from the scrounger tactic are negatively frequency dependent; a scrounger does better than a producer when the scrounger tactic is rare, but worse when it is common. No study to date has shown that the payoffs of producer and scrounger conform to the game's assumptions or that groups of foragers reach the predicted stable equilibrium frequency (SEF) of scrounger, whereby both tactics obtain the same payoff. The current study of three captive flocks of spice finches, Lonchura punctulata, provides the first test of the PS game using an apparatus in which both assumptions of the PS game are met. The payoffs to the scrounger, measured as feeding rate (seeds/s), were highly negatively frequency dependent on the frequency of scrounger. The feeding rate for scrounger declined linearly while the rate for producer either declined only slightly or not at all with increasing scrounger frequency. When given the opportunity to alternate between tactics, the birds changed their use of each, such that the group converged on the predicted SEF of scrounger after 5-8 days of testing. Individuals in this study, therefore, demonstrated sufficient plasticity in tactic use such that the flock foraged at the SEF of scrounger. Copyright 2000 The Association for the Study of Animal Behaviour.

Abstract: The way in which novel learned behaviour patterns spread through animal populations remains poorly understood, despite extensive field research and the recognition that such processes play an important role in the behavioural development, social interactions and evolution of many animal species. We conducted a series of controlled diffusions of foraging information in replicate experimental populations of the guppy, Poecilia reticulata. We presented novel foraging tasks over 15 trials to mixed-sex groups, made up of food-deprived and nonfood-deprived adults (experiment 1) or small, young fish and old, large adults (experiment 2). In these diffusions, knowledge of a route to a feeder could spread through the group by subjects learning from others, discovering the route for themselves, or, most likely, by some combination of these social and asocial learning processes. We found a striking sex difference, with novel foraging information spreading at a significantly faster rate through subgroups of females than of males. Females both discovered the goal and learned the route more quickly than males. Food-deprived individuals were faster at completing the tasks over the 15 trials than nonfood-deprived guppies, and there was a significant interaction between sex and size, with a sex difference in adults but not young individuals. There was also an interaction between sex and hunger level, with food deprivation having a stronger effect on male than female performance. We suggest that information may diffuse in a similar nonrandom or 'directed' manner through many natural populations of animals. Copyright 2000 The Association for the Study of Animal Behaviour.