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: We introduced a new technique to investigate the development of scribbling in very young infants. We tested three infant chimpanzees to compare the developmental processes of scribbling between humans and chimpanzees. While human infants start to scribble on paper at around the age of 18 months, our 13- to 23-month-old infant chimpanzees had never been observed scribbling prior to this study. We used a notebook computer with a touch-sensitive screen. This apparatus was able to record the location of the subjects' touches on the screen. Each touch generated a fingertip-sized dot at the corresponding on-screen location. During spontaneous interactions with this apparatus, all three infants and two mother chimpanzees left scribbles with their fingers on the screen. The scribbles contained not only simple dots or short lines, but also curves and hook-like lines or loops, most of which were observed in the instrumental drawings of adult chimpanzees. The results suggest that perceptual-motor control for finger drawing develops in infant chimpanzees. Two of the infants performed their first scribble with a marker on paper at the age of 20-23 months. Just prior to this, they showed a rapid increase in combinatory manipulation of objects. These findings suggest that the development of combinatory manipulation of objects as well as that of perceptual-motor control may be necessary for the emergence of instrumental drawing on paper.

Abstract: Chimpanzees (Pan troglodytes) frequently participate in social exchange involving multiple goods and services of variable value, yet they have not been tested in a formalized situation to see whether they can barter using multiple tokens and rewards. We set up a simple barter economy with two tokens and two associated rewards and tested chimpanzees on their ability to obtain rewards by returning the matching token in situations in which their access to tokens was unlimited or limited. Chimpanzees easily learned to associate value with the tokens, as expected, and did barter, but followed a simple strategy of favoring the higher-value token, regardless of the reward proffered, instead of a more complex but more effective strategy of returning the token that matched the reward. This response is similar to that shown by capuchin monkeys in our previous study. We speculate that this response, while not ideal, may be sufficient to allow for stability of the social exchange system in these primates, and that the importance of social barter to both species may have led to this convergence of strategies.

Abstract: The experiments investigated how two adult captive chimpanzees learned to navigate in an automated interception task. They had to capture a visual target that moved predictably on a touch monitor. The aim of the study was to determine the learning stages that led to an efficient strategy of intercepting the target. The chimpanzees had prior training in moving a finger on a touch monitor and were exposed to the interception task without any explicit training. With a finger the subject could move a small “ball” at any speed on the screen toward a visual target that moved at a fixed speed either back and forth in a linear path or around the edge of the screen in a rectangular pattern. Initial ball and target locations varied from trial to trial. The subjects received a small fruit reinforcement when they hit the target with the ball. The speed of target movement was increased across training stages up to 38 cm/s. Learning progressed from merely chasing the target to intercepting the target by moving the ball to a point on the screen that coincided with arrival of the target at that point. Performance improvement consisted of reduction in redundancy of the movement path and reduction in the time to target interception. Analysis of the finger's movement path showed that the subjects anticipated the target's movement even before it began to move. Thus, the subjects learned to use the target's initial resting location at trial onset as a predictive signal for where the target would later be when it began moving. During probe trials, where the target unpredictably remained stationary throughout the trial, the subjects first moved the ball in anticipation of expected target movement and then corrected the movement to steer the ball to the resting target. Anticipatory ball movement in probe trials with novel ball and target locations (tested for one subject) showed generalized interception beyond the trained ball and target locations. The experiments illustrate in a laboratory setting the development of a highly complex and adaptive motor performance that resembles navigational skills seen in natural settings where predators intercept the path of moving prey.