No abstract

Discussions of the evolution of intelligence have focused on monkeys and apes because of their close evolutionary relationship to humans. Other large-brained social animals, such as corvids, also understand their physical and social worlds. Here we review recent studies of tool manufacture, mental time travel, and social cognition in corvids, and suggest that complex cognition depends on a ''tool kit'' consisting of causal reasoning, flexibility, imagination, and prospection. Because corvids and apes share these cognitive tools, we argue that complex cognitive abilities evolved multiple times in distantly related species with vastly different brain structures in order to solve similar socioecological problems.

The ability to use tools has been suggested to indicate advanced physical cognition in animals. Here we show that rooks, a member of the corvid family that do not appear to use tools in the wild are capable of insightful problem solving related to sophisticated tool use, including spontaneously modifying and using a variety of tools, shaping hooks out of wire, and using a series of tools in a sequence to gain a reward. It is remarkable that a species that does not use tools in the wild appears to possess an understanding of tools rivaling habitual tool users such as New Caledonian crows and chimpanzees. Our findings suggest that the ability to represent tools may be a domain-general cognitive capacity rather than an adaptive specialization and questions the relationship between physical intelligence and wild tool use.cognition ͉ intelligence ͉ Corvus ͉ frugilegus ͉ hook

The ‘social intelligence hypothesis’ was originally conceived to explain how primates may have evolved their superior intellect and large brains when compared with other animals. Although some birds such as corvids may be intellectually comparable to apes, the same relationship between sociality and brain size seen in primates has not been found for birds, possibly suggesting a role for other non-social factors. But bird sociality is different from primate sociality. Most monkeys and apes form stable groups, whereas most birds are monogamous, and only form large flocks outside of the breeding season. Some birds form lifelong pair bonds and these species tend to have the largest brains relative to body size. Some of these species are known for their intellectual abilities (e.g. corvids and parrots), while others are not (e.g. geese and albatrosses). Although socio-ecological factors may explain some of the differences in brain size and intelligence between corvids/parrots and geese/albatrosses, we predict that the type and quality of the bonded relationship is also critical. Indeed, we present empirical evidence that rook and jackdaw partnerships resemble primate and dolphin alliances. Although social interactions within a pair may seem simple on the surface, we argue that cognition may play an important role in the maintenance of long-term relationships, something we name as ‘relationship intelligence’.

Social life has costs associated with competition for resources such as food. Food storing may reduce this competition as the food can be collected quickly and hidden elsewhere; however, it is a risky strategy because caches can be pilfered by others. Scrub jays (Aphelocoma coerulescens) remember 'what', 'where' and 'when' they cached. Like other corvids, they remember where conspecifics have cached, pilfering them when given the opportunity, but may also adjust their own caching strategies to minimize potential pilfering. To test this, jays were allowed to cache either in private (when the other bird's view was obscured) or while a conspecific was watching, and then recover their caches in private. Here we show that jays with prior experience of pilfering another bird's caches subsequently re-cached food in new cache sites during recovery trials, but only when they had been observed caching. Jays without pilfering experience did not, even though they had observed other jays caching. Our results suggest that jays relate information about their previous experience as a pilferer to the possibility of future stealing by another bird, and modify their caching strategy accordingly.

Western scrub-jays (Aphelocoma californica) hide food caches for future consumption, steal others' caches, and engage in tactics to minimize the chance that their own caches will be stolen. We show that scrub-jays remember which individual watched them during particular caching events and alter their recaching behavior accordingly. We found no evidence to suggest that a storer's use of cache protection tactics is cued by the observer's behavior.

The role of the amygdala in dyadic social interactions of adult rhesus monkeys (Macaca mulatta) was assessed after bilateral ibotenic acid lesions. Social, nonsocial, and spatial behaviors of amygdalectomized and control monkeys were assessed in 3 dyadic experiments: constrained, unconstrained, and round robin. Lesions produced extensive bilateral damage to the amygdala. Across all experiments, the amygdalectomized monkeys demonstrated increased social affiliation, decreased anxiety, and increased confidence compared with control monkeys, particularly during early encounters. Normal subjects also demonstrated increased social affiliation toward the amygdalectomized subjects. These results indicate that amygdala lesions in adult monkeys lead to a decrease in the species-normal reluctance to immediately engage a novel conspecific in social behavior. The altered behavior of the amygdalectomized monkeys may have induced the increased social interactions from their normal companions. This is contrary to the idea that amygdalectomy produces a decrease in social interaction and increased aggression from conspecifics.

Comparative psychologists interested in the evolution of intelligence have focused their attention on social primates, whereas birds tend to be used as models of associative learning. However, corvids and parrots, which have forebrains relatively the same size as apes, live in complex social groups and have a long developmental period before becoming independent, have demonstrated ape-like intelligence. Although, ornithologists have documented thousands of hours observing birds in their natural habitat, they have focused their attention on avian behaviour and ecology, rather than intelligence. This review discusses recent studies of avian cognition contrasting two different approaches; the anthropocentric approach and the adaptive specialization approach. It is argued that the most productive method is to combine the two approaches. This is discussed with respects to recent investigations of two supposedly unique aspects of human cognition; episodic memory and theory of mind. In reviewing the evidence for avian intelligence, corvids and parrots appear to be cognitively superior to other birds and in many cases even apes. This suggests that complex cognition has evolved in species with very different brains through a process of convergent evolution rather than shared ancestry, although the notion that birds and mammals may share common neural connectivity patterns is discussed.