Abstract: A theory of imitation is proposed, string parsing, which separates the copying of behavioural organization by observation from an understanding of the cause of its effectiveness. In string parsing, recurring patterns in the visible stream of behaviour are detected and used to build a statistical sketch of the underlying hierarchical structure. This statistical sketch may in turn aid the subsequent comprehension of cause and effect. Three cases of social learning of relatively complex skills are examined, as potential cases of imitation by string parsing. Understanding the basic requirements for successful string parsing helps to resolve the conflict between mainly negative reports of imitation in experiments and more positive evidence from natural conditions. Since string parsing does not depend on comprehension of the intentions of other agents or the everyday physics of objects, separate tests of these abilities are needed even in animals shown to learn by imitation.

Abstract: In chimpanzees, it is only in the restricted context of tool use that manual and cognitive skills have been described, comparable to those that gorillas and orang-utans display in obtaining plant foods. We report the complex food preparation skills used to eat, without tools, the leaves of the tree Broussonettia papyrifera in the Sonso community of chimpanzees at Budongo Forest, Uganda. Able-bodied individuals used multi-stage techniques that required bimanual role differentiation at several stages, and were hierarchical in organisation. A total repertoire of 14 techniques was found, with strong preference in all individuals for either of two of these; 6 additional techniques were found when flowers and leaves were eaten together. However, in this community over 20% of individuals suffer from some form of upper- or lower-limb injury as a result of snares. We investigated the manner of compensation for upper-limb injury. Only the most severely injured showed reduced feeding efficiency. Injured individuals were found to use the same repertoire of techniques as able-bodied chimpanzees. We found no evidence to suggest that injured individuals were able to develop wholly novel techniques optimal for their specific injuries, although shifts in preference for particular techniques did occur. Rather, injured individuals used novel ways of achieving particular steps in the process; by “working around” their impairments; in this way, they managed to use the same techniques as the able-bodied. Since snare injuries generally befall young animals, these results suggest that chimpanzees learn techniques partly through observational learning (of, necessarily, able-bodied individuals).

Abstract: The manipulative actions of mountain gorillas Gorilla g. beringei were examined in the context of foraging on hard-to-process plant foods in the field, in particular those used in tackling thistle Carduus nyassanus. A repertoire of 72 functionally distinct manipulative actions was recorded. Many of these actions were used in several variants of grip, finger(s) and movement path, both by different individuals and by the same individual at different times. The repertoire appears somewhat greater than that observed in comparable studies of monkeys, but a far more striking difference is found in the use of differentiated actions in concert. Mountain gorillas routinely and frequently deal with problems that involve: (1) bimanual role differentiation, with the two hands taking different roles but synchronized in time and space, and (2) digit role differentiation, with independent control of parts of the same hand used for separate purposes at the same time. The independent control that allows these abilities, so crucial to human manual constructional ability, is apparently general in African great apes. Role differentiation, between and within the hand, is evidently a primitive characteristic in the human arsenal of skills.

Abstract: Summary Underlying the various behaviors that are classified as imitation, there may be several distinct mechanisms, differing in adaptive function, cognitive basis, and computational power. Experiments reporting “true motor imitation” in animals do not as yet give evidence of production learning by imitation; instead, contextual imitation can explain their data, and this can be explained by a simple mechanism (response facilitation) which matches known neural findings. When imitation serves a function in social mimicry, which applies to a wide range of phenomena from neonatal imitation in humans and great apes to pair-bonding in some bird species, the fidelity of the behavioral match is crucial. Learning of novel behavior can potentially be achieved by matching the outcome of a model's action, and it is argued that vocal imitation by birds is a clear example of this method (which is sometimes called emulation). Alternatively, the behavior itself may be perceived in terms of actions that the observer can perform, and thus it may be copied. If the imitation is linear and stringlike (action level), following the surface form rather than the underlying plan, then its utility for learning new instrumental methods is limited. However, the underlying plan of hierarchically organized behavior is visible in output behavior, in subtle but detectable ways, and imitation could instead be based on this organization (program level), extracted automatically by string parsing. Currently, the most likely candidates for such capacities are all great apes. It is argued that this ability to perceive the underlying plan of action, in addition to allowing highly flexible imitation of novel instrumental methods, may have resulted in the competence to understand the intentions (theory of mind) of others.