Abstract: When we talk of the 'nature of intelligence', or any other attribute, we may be referring to its essential structure, or to its place in nature, particularly the function it has evolved to serve. Here I examine both, from the perspective of the evolution of intelligence in primates. Over the last 20 years, the Social (or 'Machiavellian') Intelligence Hypothesis has gained empirical support. Its core claim is that the intelligence of primates is primarily an adaptation to the special complexities of primate social life. In addition to this hypothesis about the function of intellect, a secondary claim is that the very structure of intelligence has been moulded to be 'social' in character, an idea that presents a challenge to orthodox views of intelligence as a general-purpose capacity. I shall outline the principal components of social intelligence and the environment of social complexity it engages with. This raises the question of whether domain specificity is an appropriate characterization of social intelligence and its subcomponents, like theory of mind. As a counter-argument to such specificity I consider the hypothesis that great apes exhibit a cluster of advanced cognitive abilities that rest on a shared capacity for second-order mental representation.

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: Non-human animals vary in their ability to make and use tools. The goal of the present study was to further explore what, if anything, differs between tool-users and non-tool-users, and whether these differences lie in the conceptual or motor domain. We tested two species that typically do not use tools-cotton top tamarins (Saguinus oedipus) and vervet monkeys (Cercopithecus aethiops)-on problems that mirrored those designed for prolific tool users such as chimpanzees. We trained subjects on a task in which they could choose one of two canes to obtain an out-of-reach food reward. After training, subjects received several variations on the original task, each designed to examine a specific conceptual aspect of the pulling problem previously studied in other tool-using species. Both species recognized that effective pulling tools must be made of rigid materials. Subsequent conditions revealed significant species differences, with vervets outperforming tamarins across many conditions. Vervets, but not tamarins, had some recognition of the relationship between a tool's orientation and the position of the food reward, the relationship between a tool's trajectory and the substance that it moves on, and that tools must be connected in order to work properly. These results provide further evidence that tool-use may derive from domain-general, rather than domain-specific cognitive capacities that evolved for tool use per se.

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: 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.