Abstract: Reciprocal altruism refers to the exchange of beneficial acts between individuals, in which the benefits to the recipient exceed the cost to the altruist. Theory predicts that cooperation among unrelated animals can occur whenever individuals encounter each other regularly and are capable of adjusting their cooperative behaviour according to experience. Although the potential for reciprocal altruism exists in many animal societies, most interactions occur between closely related individuals, and examples of reciprocity among non-kin are rare. The field experiments on vervet monkeys which we present here demonstrate that grooming between unrelated individuals increases the probability that they will subsequently attend to each others' solicitations for aid. Vervets appear to be more willing to aid unrelated individuals if those individuals have behaved affinitively toward them in the recent past. In contrast, recent grooming between close genetic relatives appears to have no effect on their willingness to respond to each other's solicitations for aid.

Abstract: As an increasing number of field studies of chimpanzees (Pan troglodytes) have achieved long-term status across Africa, differences in the behavioural repertoires described have become apparent that suggest there is significant cultural variation. Here we present a systematic synthesis of this information from the seven most long-term studies, which together have accumulated 151 years of chimpanzee observation. This comprehensive analysis reveals patterns of variation that are far more extensive than have previously been documented for any animal species except humans. We find that 39 different behaviour patterns, including tool usage, grooming and courtship behaviours, are customary or habitual in some communities but are absent in others where ecological explanations have been discounted. Among mammalian and avian species, cultural variation has previously been identified only for single behaviour patterns, such as the local dialects of song-birds. The extensive, multiple variations now documented for chimpanzees are thus without parallel. Moreover, the combined repertoire of these behaviour patterns in each chimpanzee community is itself highly distinctive, a phenomenon characteristic of human cultures but previously unrecognised in non-human species.

Abstract: A basic principle of human memory is that lists that can be organized into memorable 'chunks' are easier to remember. Memory span is limited to a roughly constant number of chunks and is to a large extent independent of the amount of informaton contained in each chunk. Depending on the ingenuity of the code used to integrate discrete items into chunks, one can substantially increase the number of items that can be recalled correctly. Newly developed paradigms for studying memory in non-verbal organisms allow comparison of the abilities of human and non-human subjects to memorize lists. Here I present two types of evidence that pigeons 'chunk' 5-element lists whose components (colours and achromatic geometric forms) are clustered into distinct groups. Those lists were learned twice as rapidly as a homogeneous list of colours or heterogeneous lists in which the elements are not clustered. The pigeons were also tested for knowledge of the order of two elements drawn from the 5-element lists. They responded in the correct order only to those subsets that contained a chunk boundary. Thus chunking can be studied profitably in animal subjects; the cognitive processes that allow an organism to form chunks do no presuppose linguistic competence.