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 ...

The issue whether non-human primates have other-regarding preference and/or inequity aversion has been under debate. We investigated whether tufted capuchin monkeys are sensitive to others' reward in various experimental food sharing settings. Two monkeys faced each other. The operator monkey chose one of two food containers placed between the participants, each containing a food item for him/herself and another for the recipient. The recipient passively received either high- or low-value food depending on the operator's choice, whereas the operator obtained the same food regardless of his/her choice. The recipients were either the highest- or lowest-ranking member of the group, and the operators were middle-ranking. In Experiment 1, the operators chose the high-value food for the subordinate recipient more frequently than when there was no recipient, whereas they were indifferent in their choice for the dominant. This differentiated behavior could have been because the dominant recipient frequently ate the low-value food. In Experiment 2, we increased the difference in the value of the two food items so that both recipients would reject the low-value food. The results were the same as in Experiment 1. In Experiment 3, we placed an opaque screen in front of the recipient to examine effects of visual contact between the participants. The operators' food choice generally shifted toward providing the low-value food for the recipient. These results suggest that capuchins are clearly sensitive to others' reward and that they show other-regarding preference or a form of inequity aversion depending upon the recipients and the presence of visual contact.

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Humans routinely socially evaluate others not only following direct interactions with them but also based on others' interactions with third parties. In other species, 'eavesdropping' on third-party interactions is often used to gain information about foraging or mating opportunities, or others individuals' aggressiveness or fighting ability. However, image scoring for potential cooperativeness is less well studied. Here we ask whether a non-human primate species, tufted capuchin monkeys (Cebus apella), socially evaluates humans after witnessing third-party interactions involving a helpful intervention or failure to help. We find that the monkeys accept food less frequently from those who persistently reject another's requests for help. This negative social evaluation effect is robust across conditions, and tightly linked to explicit refusal to help. Evaluation of potential helpfulness based on third-party interactions may thus not be unique to humans.

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Our society is sustained by wide-ranging cooperation. If individuals are sensitive to others' gains and losses as well as the amount of labor, they can ensure future beneficial cooperative interaction. However, it is still an open question whether nonhuman primates are sensitive to others' labor. We asked this question in tufted capuchin monkeys in an experimental food-sharing situation by comparing conditions with labor by two participants equalized (Equal labor condition) or unequalized (Unequal labor condition). The operator monkey pulled the drawer of one of the two food containers placed between two monkeys, each containing a food for him/herself and another for the recipient monkey. The recipient received either high- or low-value food depending on the operator's choice, whereas the operator obtained the same food regardless of his/her choice. In Unequal labor condition, the operator first had to pull the handle of the board to which the containers were glued and then pull the drawer of one of the containers, while the recipient received food with no labor. In Equal labor condition, the recipient had to pull the handle of the board so that the operator could operate a container. Results showed that operators chose the high-value food container for recipients more often than when the recipient was absent only in Equal labor condition. This suggests that capuchin monkeys are sensitive to others' labor and actively give food to a partner who has helped them to complete a task.

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We tested whether horses (Equus caballus) are sensitive to human attentional states and modify the modality of begging behaviors as a function of human attentional states in a naturalistic food-requesting situation. In Experiment 1, horses tended to produce more auditory or tactile begging behaviors when the human experimenter (E1)'s eyes were covered by her hand than when they were not covered. However, there was no difference in visual begging behaviors between conditions. In Experiment 2, horses produced significantly more auditory or tactile begging behaviors when E1's eyes were closed than when they were open. In contrast, horses produced significantly more visual begging behaviors when E1's eyes were open than when they were closed. These results suggest that horses understand the role of eyes as an indicator of human attentional states and show effective and flexible begging behaviors proactively as a function of human attentional states. ey oodss: K horses, human attentional states, begging behaviors