Abstract: Great ape gesture has become a research topic of intense interest, because its intentionality and flexibility suggest strong parallels to human communication. Yet the fundamental question of whether an animal species’ gestures carry specific meanings has hardly been addressed. We set out a systematic approach to studying intentional meaning in the gestural communication of non-humans and apply it to a sample of orangutan gestures. We propose that analysis of meaning should be limited to gestures for which (1) there is strong evidence for intentional production and (2) the recipient’s final reaction matches the presumed goal of the signaller, as determined independently. This produces a set of successful instances of gesture use, which we describe as having goal–outcome matches. In this study, 28 orangutans in three European zoos were observed for 9 months. We distinguished 64 gestures on structural grounds, 40 of which had frequent goal–outcome matches and could therefore be analysed for meaning. These 40 gestures were used predictably to achieve one of 6 social goals: to initiate an affiliative interaction (contact, grooming, or play), request objects, share objects, instigate co-locomotion, cause the partner to move back, or stop an action. Twenty-nine of these gestures were used consistently with a single meaning. We tested our analysis of gesture meaning by examining what gesturers did when the response to their gesture did not match the gesture’s meaning. Subsequent actions of the gesturer were consistent with our assignments of meaning to gestures. We suggest that, despite their contextual flexibility, orangutan gestures are made with the expectation of specific behavioural responses and thus have intentional meanings as well as functional consequences.

Abstract: A growing body of evidence suggests that a wide range of animals can recognize and respond appropriately to calls produced by other species. Social learning has been implicated as a possible mechanism by which heterospecific call recognition might develop. To examine whether familiarity and/or shared vulnerability with the calling species might influence the ability of sympatric species to distinguish heterospecific alarm calls, we tested whether four ungulate species (impala: Aepyceros melampus; tsessebe: Damaliscus lunatus; zebra: Equus burchelli; wildebeest: Connochaetes taurinus) could distinguish baboon (Papio hamadryas ursinus) alarm calls from other loud baboon calls produced during intra-specific aggressive interactions (‘contest’ calls). Overall, subjects’ responses were stronger following playback of alarm calls than contest calls. Of the species tested, impala showed the strongest responses and the greatest difference in composite response scores, suggesting they were best able to differentiate call types. Compared with the other ungulate species, impala are the most frequent associates of baboons. Moreover, like baboons, they are susceptible to both lion and leopard attacks, whereas leopards rarely take the larger ungulates. Although it seems possible that high rates of association and/or shared vulnerability may influence impala’s greater ability to distinguish among baboon call types, our results point to a stronger influence of familiarity. Ours is the first study to compare such abilities among several community members with variable natural histories, and we discuss future experiments that would more systematically examine development of these skills in young ungulates.

Abstract: Demonstrating the ability to ‘copy’ the behavior of others is an important aspect in determining whether social learning occurs and whether group level differences in a given behavior represent cultural differences or not. Understanding the occurrence of this process in its natural context is essential, but can be a daunting task in the wild. In order to test the social learning hypothesis for the acquisition of leaf swallowing (LS), a self-medicative behavior associated with the expulsion of parasites, we conducted semi-naturalistic experiments on two captive groups of parasite-free, naïve chimpanzees (Pan troglodytes). Individuals in the group were systematically provided appropriate stimuli (rough hispid leaves) identical to those used by chimpanzees in the wild. Individuals initially responded in a variety of ways, ranging from total aversion to normal chewing and swallowing. Over time, however, the two groups adopted different variants for inserting and folding the leaves in the mouth prior to swallowing them (complete and partial LS), following the specific method spontaneously displayed by the first and primary LS models in their respective groups. These variants were similar to LS displayed by chimpanzees in the wild. Using the option-bias method, we found evidence for social learning leading to group-level biased transmission and group-level stabilization of these two variants. This is the first report on two distinct cultural variants innovated in response to the introduction of natural stimuli that emerged and spread spontaneously and concurrently within two adjacent groups of socially housed primates. These observations support the assertion that LS may reflect a generalized propensity for ingesting rough hispid leaves, which can be socially induced and transmitted within a group. Ingesting an adequate number of these leaves induces increased gut motility, which is responsible for the subsequent expulsion of particular parasite species in the wild. Cultural transmission and maintenance of LS within a group and associative learning by the individual of the positive consequences of this otherwise non-nutritive mode of ingestion is proposed to be the pivotal link between this feeding propensity and its maintenance as a self-medicative behavior by great apes in the wild.

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: Selection pressures during domestication are thought to lead to an enhanced ability to use human-given cues. Horses fulfil a wide variety of roles for humans and have been domesticated for at least 5000 years but their ability to read human cues has not been widely studied. We tested the ability of 28 horses to attend to human-given cues in an object choice task. We included five different cues: distal sustained pointing, momentary tapping, marker placement, body orientation and gaze (head) alternation. Horses were able to use the pointing and marker placement cues spontaneously but not the tapping, body orientation and gaze alternation cues. The overall pattern of responding suggests that horses may use cues that provide stimulus enhancement at the time of choice and do not have an understanding of the communicative nature of the cues given. As such, their proficiency at this task appears to be inferior to that of domestic dogs, Canis lupus familiaris, but similar to that of domestic goats, Caprus hircus.