Cognition presents evolutionary research with one of its greatest challenges. Cognitive evolution has been explained at the proximate level by shifts in absolute and relative brain volume and at the ultimate level by differences in social and dietary complexity. However, no study has integrated the experimental and phylogenetic approach at the scale required to rigorously test these explanations. Instead, previous research has largely relied on various measures of brain size as proxies for cognitive abilities. We experimentally evaluated these major evolutionary explanations by quantitatively comparing the cognitive performance of 567 individuals representing 36 species on two problem-solving tasks measuring self-control. Phylogenetic analysis revealed that absolute brain volume best predicted performance across species and accounted for considerably more variance than brain volume controlling for body mass. This result corroborates recent advances in evolutionary neurobiology and illustrates the cognitive consequences of cortical reorganization through increases in brain volume. Within primates, dietary breadth but not social group size was a strong predictor of species differences in self-control. Our results implicate robust evolutionary relationships between dietary breadth, absolute brain volume, and self-control. These findings provide a significant first step toward quantifying the primate cognitive phenome and explaining the process of cognitive evolution.psychology | behavior | comparative methods | inhibitory control | executive function S ince Darwin, understanding the evolution of cognition has been widely regarded as one of the greatest challenges for evolutionary research (1). Although researchers have identified surprising cognitive flexibility in a range of species (2-40) and potentially derived features of human psychology (41-61), we know much less about the major forces shaping cognitive evolution (62-71). With the notable exception of Bitterman's landmark studies conducted several decades ago (63, 72-74), most research comparing cognition across species has been limited to small taxonomic samples (70, 75). With limited comparable experimental data on how cognition varies across species, previous research has largely relied on proxies for cognition (e.g., brain size) or metaanalyses when testing hypotheses about cognitive evolution (76-92). The lack of cognitive data collected with similar methods across large samples of species precludes meaningful species comparisons that can reveal the major forces shaping cognitive evolution across species, including humans (48,70,89,(93)(94)(95)(96)(97)(98). SignificanceAlthough scientists have identified surprising cognitive flexibility in animals and potentially unique features of human psychology, we know less about the selective forces that favor cognitive evolution, or the proximate biological mechanisms underlying this process. We tested 36 species in two problemsolving tasks measuring self-control and evaluated the leading hypotheses regarding how ...

Recent research with Rooks has demonstrated impressive tool-using abilities in captivity despite this species' classification as a non-tool-user in the wild. Here, we explored whether another non-tool-using corvid, the Eurasian Jay, would be capable of similar feats and investigated the relative contributions of causal knowledge and instrumental conditioning to the birds' performance on the tasks. Five jays were tested on a variety of tasks involving water displacement. Two birds reliably interacted with the apparatuses. In these tasks, both birds showed a preference for inserting stones into a tube containing liquid over a tube containing a solid or a baited 'empty' tube and also for inserting sinkable items over non-sinkable items into a tube of water. To investigate the contribution of instrumental conditioning, subjects were then tested on a series of tasks in which different cues were made available. It was found that, in the absence of any apparent causal cues, these birds showed a clear preference for the rewarded tube when the food incrementally approached with every stone insertion, but not when it simply "appeared" after the correct number of stone insertions. However, it was found that subjects did not prefer to insert stones into a tube rewarded by the incremental approach of food if the available causal cues violated the expectations created by existing causal knowledge (i.e. were counter-intuitive). An analysis of the proportion of correct and incorrect stone insertions made in each trial across tasks offering different types of information revealed that subjects were substantially more successful in experiments in which causal cues were available, but that rate of learning was comparable in all experiments. We suggest that these results indicate that Eurasian jays use the incremental approach of the food reward as a conditioned reinforcer allowing them to solve tasks involving raising the water level and that this learning is facilitated by the presence of causal cues.

Understanding causal regularities in the world is a key feature of human cognition. However, the extent to which non-human animals are capable of causal understanding is not well understood. Here, we used the Aesop's fable paradigm – in which subjects drop stones into water to raise the water level and obtain an out of reach reward – to assess New Caledonian crows' causal understanding of water displacement. We found that crows preferentially dropped stones into a water-filled tube instead of a sand-filled tube; they dropped sinking objects rather than floating objects; solid objects rather than hollow objects, and they dropped objects into a tube with a high water level rather than a low one. However, they failed two more challenging tasks which required them to attend to the width of the tube, and to counter-intuitive causal cues in a U-shaped apparatus. Our results indicate that New Caledonian crows possess a sophisticated, but incomplete, understanding of the causal properties of displacement, rivalling that of 5–7 year old children.

State-attribution is the ability to ascribe to others an internal life like one's own and to understand that internal, psychological states such as desire, hope, belief, and knowledge underlie others' actions. Despite extensive research, comparative studies struggle to adequately integrate key factors of state-attribution that have been identified by evolutionary and developmental psychology as well as research on empathy. Here, we develop a behavioral paradigm to address these issues and investigate whether male Eurasian jays respond to the changing desire-state of their female partners when sharing food. We demonstrate that males feed their mates flexibly according to the female's current food preference. Critically, we show that the males need to see what the female has previously eaten to know what food she will currently want. Consequently, the males' sharing pattern was not simply a response to their mate's behavior indicating her preference as to what he should share, nor was it a response to the males' own desire-state. Our results raise the possibility that these birds may be capable of ascribing desire to their mates.corvid | Theory of Mind | cooperation | specific satiety I n human development, the earliest manifestation of state-attribution is the ability to explain and predict others' behavior in terms of desire-states (e.g., wants, wishes, hopes), a capacity that does not require a concept of others' mental representations (1, 2). At around the age of four, the more complex desire-belief system is formed by incorporating the attribution of epistemic mental states (e.g., belief, knowledge, doubt, expectation) (3, 4), an ability known as Theory of Mind. It is very possible that the attribution of desire-states may represent not only the developmental, but also the evolutionary, precursor of Theory of Mind. Thus, the first logical step in the investigation of state-attribution in nonhuman animals should be to search for this simpler manifestation (5). Despite this, most comparative studies focus solely on the attribution of epistemic mental states (6-13).A notable exception claims that apes can retrospectively infer humans' desires toward food items from the emotions expressed (14). However, given that the apes were explicitly trained about the relationship between human expressions and the availability of food, we cannot conclude that the apes inferred an underlying desire. In another study, the authors claimed that capuchin monkeys (Cebus apella) attribute a need or desire to another individual and thus might exhibit empathetic perspective-taking because the number of tolerated thefts by another individual was lower after subjects had seen the other individual eat than when they had not (15). Unfortunately, this study failed to account for a much simpler competitive account, namely that the sight of another monkey eating triggered the monkeys to protect their food. A second issue is that this experimental paradigm could only investigate the understanding of another individual's physiological process...

Studies on members of the crow family using the “Aesop's Fable” paradigm have revealed remarkable abilities in these birds, and suggested a mechanism by which associative learning and folk physics may interact when learning new problems. In the present study, children between 4 and 10 years of age were tested on the same tasks as the birds. Overall the performance of the children between 5–7-years was similar to that of the birds, while children from 8-years were able to succeed in all tasks from the first trial. However the pattern of performance across tasks suggested that different learning mechanisms might be being employed by children than by adult birds. Specifically, it is possible that in children, unlike corvids, performance is not affected by counter-intuitive mechanism cues.