Observer choices during experimental foraging tasks in brown capuchin monkeys (Cebus apella).

We investigated whether capuchin monkeys (Cebus apella) would choose to observe a high- or low-status adult female from their group during experimental foraging tests. The subject was located in the center of a test chamber, with a low- and high-ranking demonstrator on either side of two partitions. A peephole allowed the subject to observe the models by looking through either respective partition. Each model was trained on one of the two different methods, lift or pull, for retrieving food from a foraging apparatus. There were 22 subjects and four models. During the 40-trial test sessions, subjects could choose which model they would watch in each trial. It was predicted that subjects would prefer observing the model with whom it was closer in rank, and therefore share greater affiliation with. Results showed that only half the subjects showed a preference and that preference was not linked to status. Relatedness played a larger role in determining if a subject showed a preference for a model, and a correlation was found for relatedness and observer preference. After the observer preference tests, subjects were presented with the foraging apparatus to determine if they displayed a preference for one of the two tasks. The majority of subjects (17/22) showed a preference for the pull method, suggesting that this method may have been more salient to the monkeys in this study.

Observational learning in orangutan cultural transmission chains.

Field reports suggest that orangutans acquire local traditions by observing neighbouring conspecifics. However, there is little direct evidence of social learning to support this conclusion. The present study investigated whether orangutans would learn a novel foraging method through observation of a conspecific in a diffusion-chain paradigm testing for the spread of the behaviour. A foraging box with two possible methods for extracting food was used to investigate the transmission of a foraging tradition among zoo-living subjects. In a socially housed group of five orangutans, the dominant male was trained to use one technique exclusively to retrieve food. He then performed this technique in the presence of another group member, who was then allowed to forage. After several trials, the observer became the model for the next individual. A second experimental group of six individuals was introduced to the alternative method. The model-seeded technique was successfully transmitted along both experimental chains, with significant preferences for the model method. These results are consistent with claims for social transmission of foraging methods in wild orangutans.

Conditional copying fidelity in capuchin monkeys (Cebus apella).

In the last two decades, it became largely accepted that monkeys show little, if any, copying fidelity. However, some recent studies have begun to challenge this notion. To explore reasons for such contrary findings, we designed a foraging apparatus so that in each of two experiments with capuchin monkeys (Cebus apella), a model would demonstrate one of two alternative methods to obtain food. The apparatus had a V-shaped track on which a panel could be slid up left or right from the center to reveal food. In Experiment 1, food was located in a cup directly behind the center panel. In Experiment 2, sliding the panel left or right revealed food either in left or right ends of the V-track. Since this sliding movement led directly to one food location exclusive of the other, we predicted capuchins would show greater copying fidelity in this second Experiment. Instead, subjects were significantly more faithful to the model's method in Experiment 1, which provided strong evidence of capuchins copying what they had observed. We suggest that the contrasting results of Experiment 1 may have occurred because capuchins prioritize exploratory behavior when alternative foraging locations are accessible.

In-group conformity sustains different foraging traditions in capuchin monkeys (Cebus apella).

BACKGROUND: Decades of research have revealed rich cultural repertoires encompassing multiple traditions in wild great apes, a picture crucially complemented by experimental simulations with captive apes. Studies with wild capuchin monkeys, the most encephalized simian species, have indicated a New World convergence on these cultural phenomena, involving multiple traditions and tool use. However, experimental studies to date are in conflict with such findings in concluding that capuchins, like other monkeys, show minimal capacities for social learning. METHODOLOGY/PRINCIPAL FINDINGS: Here we report a new experimental approach in which the alpha male of each of two groups of capuchins was trained to open an artificial foraging device in a quite different way, using either a slide or lift action, then reunited with his group. In each group a majority of monkeys, 8 of 11 and 13 of 14, subsequently mastered the task. Seventeen of the successful 21 monkeys discovered the alternative action to that seeded in the group, performing it a median of 4 times. Nevertheless, all 21 primarily adopted the technique seeded by their group's alpha male. Median proportions of slide versus lift were 0.96 for the group seeded with slide versus 0. 01 for the group seeded with lift. CONCLUSIONS/SIGNIFICANCE: These results suggest a striking effect of social conformity in learned behavioral techniques, consistent with field reports of capuchin traditions and convergent on the only other species in which such cultural phenomena have been reported, chimpanzees and humans.

Social facilitation of exploratory foraging behavior in capuchin monkeys (Cebus apella).

Much of the research on animal social learning focuses on complex cognitive functions such as imitation and emulation. When compelling evidence for such processes is not forthcoming, simpler processes are often assumed but rarely directly tested for. In this study we address the phenomenon of social facilitation, whereby the presence of a feeding conspecific is hypothesized to affect the motivation and behavior of the subject, elevating the likelihood of exploration and discovery in relation to the task at hand. Using a novel foraging task, sufficiently challenging that only just over half the subjects successfully gained food from it, we compared the performance of capuchin monkeys working either alone, or in a "social" condition where an actively feeding conspecific was in an adjacent chamber. Although similar numbers of subjects in these conditions were eventually successful during the 20 trials presented, the latency to successful solution of the task was over three times faster for monkeys in the social condition. The minority of monkeys that failed to learn (9/23) were then exposed to a proficient model. Only those older than 5 years provided evidence of learning from this. Accordingly, we obtained evidence for the social facilitation the study was designed to test for, and limited supplementary evidence for social learning in the older individuals who had not learned individually. These results are discussed in relation to other recent evidence for social learning in monkeys.

Social diffusion of novel foraging methods in brown capuchin monkeys (Cebus apella).

It has been reported that wild capuchin monkeys exhibit several group-specific behavioural traditions. By contrast, experiments have found little evidence for the social learning assumed necessary to support such traditions. The present study used a diffusion chain paradigm to investigate whether a novel foraging task could be observationally learned by capuchins (Cebus apella) and then transmitted along a chain of individuals. We used a two-action paradigm to control for independent learning. Either of two methods (lift or slide) could be used to open the door of a foraging apparatus to retrieve food. Two chains were tested (N1=4; N2=5), each beginning with an experimenter-trained model who demonstrated to a partner its group-specific method for opening the foraging apparatus. After the demonstration, if the observer was able to open the apparatus 20 times by either method, then it became the demonstrator for a new subject, thus simulating the spread of a foraging tradition among 'generations' of group members. Each method was transmitted along these respective chains with high fidelity, echoing similar results presently available only for chimpanzees and children. These results provide the first clear evidence for faithful diffusion of alternative foraging methods in monkeys, consistent with claims for capuchin traditions in the wild.

Partner effects on food consumption in brown capuchin monkeys.

It has been claimed that capuchin monkeys (Cebus apella) show inequity aversion in relation to food rewards for a simple exchange task. However, other factors may affect the willingness of a monkey to consume foods of high or low value in the presence of a conspecific. In this study, pairs of monkeys were presented with unequally valued foods, but without any task-performance: they simply received the food under four experimental conditions. By looking at the rate of collection and consumption of low-valued cucumber slices we expected to see variation dependent on whether the partner either had 1) cucumber (equity), 2) grape (inequity), 3) inaccessible cucumber or 4) inaccessible grape. Testing 12 adult capuchin monkeys, our findings differed from those of other authors in that the monkeys failed to show negative reactions to inequity, but rather responded with scramble competition (i.e., fast food collection) in the presence of a conspecific without access to food. They also showed facilitated consumption in the presence of a conspecific consuming high-valued food. Possibly, (in)equity plays a different role if food serves as a reward for a task rather than if it is simply made available for consumption.

The monkey in the mirror: hardly a stranger.

It is widely assumed that monkeys see a stranger in the mirror, whereas apes and humans recognize themselves. In this study, we question the former assumption by using a detailed comparison of how monkeys respond to mirrors versus live individuals. Eight adult female and six adult male brown capuchin monkeys (Cebus apella) were exposed twice to three conditions: (i) a familiar same-sex partner, (ii) an unfamiliar same-sex partner, and (iii) a mirror. Females showed more eye contact and friendly behavior and fewer signs of anxiety in front of a mirror than they did when exposed to an unfamiliar partner. Males showed greater ambiguity, but they too reacted differently to mirrors and strangers. Discrimination between conditions was immediate, and blind coders were able to tell the difference between monkeys under the three conditions. Capuchins thus seem to recognize their reflection in the mirror as special, and they may not confuse it with an actual conspecific. Possibly, they reach a level of self-other distinction intermediate between seeing their mirror image as other and recognizing it as self.