Abstract: The transmission of tool use is a rare event in monkeys. Such an event arose in a group of semi-free-ranging Tonkean macaques (Macaca tonkeana) in which leaning a pole against the park's fence (branch leaning) appeared and spread to several males. This prompted us to test individual and social learning of this behavior in seven young males. In the first experiment, three males learned individually to obtain a food reward using a wooden pole as a climbing tool. They began using the pole to retrieve the reward only when they could alternatively experience acting on the object and reaching the target. In a second experiment, we first tested whether four other subjects could learn branch leaning after having observed a group-mate performing the task. Despite repeated opportunities to observe the demonstrator, they did not learn to use the pole as a tool. Hence we exposed the latter subjects to individual learning trials and they succeeded in the task. Tool use was not transmitted in the experimental situation, which contrasts with observations in the park. We can conclude that the subjects were not able to recognize the target as such. It is possible that they recognized it and learned the task individually when we alternated the opportunity to act upon the object and to reach the reward. This suggests that these macaques could then have associated the action they exercised upon the pole and the use of the pole as a means to reach the reward.

Abstract: The ability to act on information flexibly is one of the cornerstones of intelligent behavior. As particularly informative example, tool-oriented behavior has been investigated to determine to which extent nonhuman animals understand means-end relations, object affordances, and have specific motor skills. Even planning with foresight, goal-directed problem solving and immediate causal inference have been a focus of research. However, these cognitive abilities may not be restricted to tool-using animals but may be found also in animals that show high levels of curiosity, object exploration and manipulation, and extractive foraging behavior. The kea, a New Zealand parrot, is a particularly good example. We here review findings from laboratory experiments and field observations of keas revealing surprising cognitive capacities in the physical domain. In an experiment with captive keas, the success rate of individuals that were allowed to observe a trained conspecific was significantly higher than that of naive control subjects due to their acquisition of some functional understanding of the task through observation. In a further experiment using the string-pulling task, a well-probed test for means-end comprehension, we found the keas finding an immediate solution that could not be improved upon in nine further trials. We interpreted their performance as insightful in the sense of being sensitive of the relevant functional properties of the task and thereby producing a new adaptive response without trial-and-error learning. Together, these findings contribute to the ongoing debate on the distribution of higher cognitive skills in the animal kingdom by showing high levels of sensorimotor intelligence in animals that do not use tools. In conclusion, we suggest that the 'Technical intelligence hypothesis' (Byrne, Machiavellian intelligence II: extensions and evaluations, pp 289-211, 1997), which has been proposed to explain the origin of the ape/monkey grade-shift in intelligence by a selection pressure upon an increased efficiency in foraging behavior, should be extended, that is, applied to some birds as well.

Abstract: To date, neither primates nor birds have shown clear evidence of causal knowledge when attempting to solve the trap tube task. One factor that may have contributed to mask the knowledge that subjects may have about the task is that subjects were only allowed to push the reward away from them, which is a particularly difficult action for primates in certain problem solving situations. We presented five orangutans (Pongo pygmaeus), two chimpanzees (Pan troglodytes), two bonobos (Pan paniscus), and one gorilla (Gorilla gorilla) with a modified trap tube that allowed subjects to push or rake the reward with the tool. In two additional follow-up tests, we inverted the tube 180 degrees rendering the trap nonfunctional and also presented subjects with the original task in which they were required to push the reward out of the tube. Results showed that all but one of the subjects preferred to rake the reward. Two orangutans and one chimpanzee (all of whom preferred to rake the reward), consistently avoided the trap only when it was functional but failed the original task. These findings suggest that some great apes may have some causal knowledge about the trap-tube task. Their success, however, depended on whether they were allowed to choose certain tool-using actions.

Abstract: Humans, chimpanzees, capuchins and aye-ayes all display an unusually high degree of encephalization and diverse omnivorous extractive foraging. It has been suggested that the high degree of encephalization in aye-ayes may be the result of their diverse, omnivorous extractive foraging behaviors. In combination with certain forms of tool use, omnivorous extractive foraging has been hypothesized to be linked to higher levels of sensorimotor intelligence (stages 5 or 6). Although free-ranging aye-ayes have not been observed to use tools directly in the context of their extractive foraging activities, they have recently been reported to use lianas as tools in a manner that independently suggests that they may possess stage 5 or 6 sensorimotor intelligence. Although other primate species which display diverse, omnivorous extractive foraging have been tested for sensorimotor intelligence, aye-ayes have not. We report a test of captive aye-ayes' comprehension of tool use in a situation designed to simulate natural conditions. The results support the view that aye-ayes do not achieve stage 6 comprehension of tool use, but rather may use trial-and-error learning to develop tool-use behaviors. Other theories for aye-aye encephalization are considered.

Abstract: Despite considerable current interest in the evolution of intelligence, the intuitively appealing notion that brain volume and “intelligence” are linked remains untested. Here, we use ecologically relevant measures of cognitive ability, the reported incidence of behavioral innovation, social learning, and tool use, to show that brain size and cognitive capacity are indeed correlated. A comparative analysis of 533 instances of innovation, 445 observations of social learning, and 607 episodes of tool use established that social learning, innovation, and tool use frequencies are positively correlated with species' relative and absolute “executive” brain volumes, after controlling for phylogeny and research effort. Moreover, innovation and social learning frequencies covary across species, in conflict with the view that there is an evolutionary tradeoff between reliance on individual experience and social cues. These findings provide an empirical link between behavioral innovation, social learning capacities, and brain size in mammals. The ability to learn from others, invent new behaviors, and use tools may have played pivotal roles in primate brain evolution.