Abstract: We review socially biased learning about food and problem solving in monkeys, relying especially on studies with tufted capuchin monkeys (Cebus apella) and callitrichid monkeys. Capuchin monkeys most effectively learn to solve a new problem when they can act jointly with an experienced partner in a socially tolerant setting and when the problem can be solved by direct action on an object or substrate, but they do not learn by imitation. Capuchin monkeys are motivated to eat foods, whether familiar or novel, when they are with others that are eating, regardless of what the others are eating. Thus, social bias in learning about foods is indirect and mediated by facilitation of feeding. In most respects, social biases in learning are similar in capuchins and callitrichids, except that callitrichids provide more specific behavioral cues to others about the availability and palatability of foods. Callitrichids generally are more tolerant toward group members and coordinate their activity in space and time more closely than capuchins do. These characteristics support stronger social biases in learning in callitrichids than in capuchins in some situations. On the other hand, callitrichids' more limited range of manipulative behaviors, greater neophobia, and greater sensitivity to the risk of predation restricts what these monkeys learn in comparison with capuchins. We suggest that socially biased learning is always the collective outcome of interacting physical, social, and individual factors, and that differences across populations and species in social bias in learning reflect variations in all these dimensions. Progress in understanding socially biased learning in nonhuman species will be aided by the development of appropriately detailed models of the richly interconnected processes affecting learning.

Abstract: The behaviour of members of a group of rhesus monkeys was observed in experimentally created, competitive feeding situations. Socially dominant members of the group tended to start eating before lower-ranking subjects, and generally ate more. Dominants sometimes used aggression to control access to food, but overall, intermediate-ranking monkeys were involved in most agonistic episodes. Non-dominant subjects improved their feeding performance when food was presented in three piles rather than one pile, often by snatching food and consuming it away from the pile. These general patterns were less evident when realistic snake models were placed on some of the food piles. Feeding was disrupted by the presence of snakes, but notably, subordinates risked feeding in these conditions. Piles containing preferred foods and snakes were eaten from, but a low-preference food (carrot) under snakes went untouched by all subjects. The results suggest that group-members evaluate potential risks and benefits of competing for a restricted resource, and that dominance status, while an important factor, is only one element in the equation.

Abstract: An animal's within-group spatial position has several important fitness consequences. Risk of predation, time spent engaging in antipredatory behavior and feeding competition can all vary with respect to spatial position. Previous research has found evidence that feeding rates are higher at the group edge in many species, but these studies have not represented the entire breadth of dietary diversity and ecological situations faced by many animals. In particular the presence of concentrated, defendable food patches can lead to increased feeding rates by dominants in the center of the group that are able to monopolize or defend these areas. To fully understand the tradeoffs of within-group spatial position in relation to a variety of factors, it is important to be able to predict where individuals should preferably position themselves in relation to feeding rates and food competition. A qualitative model is presented here to predict how food depletion time, abundance of food patches within a group, and the presence of prior knowledge of feeding sites affect the payoffs of different within-group spatial positions for dominant and subordinate animals. In general, when feeding on small abundant food items, individuals at the front edge of the group should have higher foraging success. When feeding on slowly depleted, rare food items, dominants will often have the highest feeding rates in the center of the group. Between these two extreme points of a continuum, an individual's optimal spatial position is predicted to be influenced by an additional combination of factors, such as group size, group spread, satiation rates, and the presence of producer-scrounger tactics.

Abstract: We tested for social learning and imitation in common marmosets using an artificial foraging task and trained conspecific demonstrators. We trained a demonstrator marmoset to open an artificial fruit, providing a full demonstration of the task to be learned. Another marmoset provided a partial demonstration, controlling for stimulus enhancement effects, by eating food from the outside of the apparatus. We thus compared three observer groups, each consisting of four animals: those that received the full demonstration, those that received the partial demonstration, and a control group that saw no demonstration prior to testing. Although none of the observer marmosets succeeded in opening the artificial fruit during the test periods, there were clear effects of demonstration type. Those that saw the full demonstration manipulated the apparatus more overall, whereas those from the control group manipulated it the least of the three groups. Those from the full-demonstration group also contacted the particular parts of the artificial fruit that they had seen touched (localised stimulus enhancement) to a greater extent than the other two groups. There was also an interaction between the number of hand and mouth touches made to the artificial fruit for the full- and partial-demonstration groups. Whether or not these data represent evidence for imitation is discussed. We also propose that the clear differences between the groups suggest that social learning mechanisms provide real benefits to these animals in terms of developing novel food-processing skills analogous to the one presented here.

Abstract: Although field studies have suggested the existence of cultural transmission of foraging techniques in primates, identification of transmission mechanisms has remained elusive. To test experimentally for evidence of imitation in the current study, we exposed gorillas (Gorilla gorilla gorilla) to an artificial fruit foraging task designed by A. Whiten and D. M. Custance (1996). Gorillas (n=6) watched a human model remove a series of 3 defenses around a fruit. Each of the defenses was removed using 1 of 2 alternative techniques. Subsequent video analysis of gorillas' behavior showed a significant tendency to copy the observed technique on 1 of the individual defenses and the direction of removal on another defense. This is the first statistically reliable evidence of imitation in gorillas. Sequence of defense removal was not replicated. The gorillas' responses were most similar to those of chimpanzees.