Calculated reciprocity after all: computation behind token transfers in orang-utans.

Transfers and services are frequent in the animal kingdom. However, there is no clear evidence in animals that such transactions are based on weighing costs and benefits when giving or returning favours and keeping track of them over time (i.e. calculated reciprocity). We tested two orang-utans (Pongo pygmaeus abelii) in a token-exchange paradigm, in which each individual could exchange a token for food with the experimenter but only after first obtaining the token from the other orang-utan. Each orang-utan possessed tokens valuable to their partner but useless to themselves. Both orang-utans actively transferred numerous tokens (mostly partner-valuable) to their partner. One of the orang-utans routinely used gestures to request tokens while the other complied with such requests. Although initially the transfers were biased in one direction, they became more balanced towards the end of the study. Indeed, data on the last three series produced evidence of reciprocity both between and within trials. We observed an increase in the number and complexity of exchanges and alternations. This study is the first experimental demonstration of the occurrence of direct transfers of goods based on calculated reciprocity in non-human-primates.

Chimpanzee (Pan troglodytes) anticipation of food return: coping with waiting time in an exchange task.

Despite controversial expectations that animals achieve reciprocal altruism, it is unclear if nonhuman species possess the necessary cognitive abilities. For reciprocal altruism, individuals must anticipate the loss of a commodity and accept a delay before some return. The authors investigated the abilities of 5 chimpanzees (Pan troglodytes) to cope with increasing waiting duration in exchange tasks. Subjects had to keep a small cookie before returning it to a human partner to obtain a larger piece. For a piece 2, 4, or 8 times the size of the small piece, 3 of the 5 subjects waited for up to 4 min. For a piece 40 times larger, 4 of the 5 subjects waited up to 8 min. At long time lag, renouncement to wait occurred earlier than predicted by subjects' general waiting capacity, suggesting that the decision to wait was based on a trade-off between reward quantity and expected costs of the waiting duration. Chimpanzees could anticipate a delayed reward at a time scale of several minutes. If this reflects a cognitive limit in chimpanzees' anticipation capacity, reciprocal altruism by keeping track of costs and benefits over extended periods may be unlikely in chimpanzees.

Accepting loss: the temporal limits of reciprocity in brown capuchin monkeys.

Delayed reciprocity is a potentially important mechanism for cooperation to occur. It is however rarely reported among animals, possibly because it requires special skills like the ability to plan a loss. We tested six brown capuchin monkeys (Cebus apella) in such skills. Subjects were studied in exchange tasks in which they had to retain a food item for a given time lag before returning it to an experimenter and obtaining a more desirable reward. Experiments showed that the subjects could wait for several minutes when allowed to return only part of the initial item. When required to return the full item intact, however, most subjects could not sustain a time lag longer than 10 s. Although the duration of waiting increased with the amount of return expected by subjects, in most cases it did not extend beyond 20 s even when the eperimenter offered a food amount 40 fold the initial item. The failure of capuchin monkeys to sustain long-lasting waiting periods may be explained by limited self-control abilities. This would prevent them achieving reciprocal altruism.