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 ...

Despite sharing a recent common ancestor, humans are surprisingly different from other great apes. The most obvious discontinuities are related to our cognitive abilities, including language, but we also have a markedly different, cooperative breeding system. Among many nonhuman primates and mammals in general, cooperative breeding is accompanied by psychological changes leading to greater prosociality, which directly enhances performance in social cognition. Here we propose that these cognitive consequences of cooperative breeding could have become more pervasive in the human lineage because the psychological changes were added to an ape-level cognitive system capable of understanding simple mental states, albeit mainly in competitive contexts. Once more prosocial motivations were added, these cognitive abilities could also be used for cooperative purposes, including a willingness to share mental states, thereby enabling the emergence of shared intentionality. Shared intentionality has been identified as the original source of many uniquely human cognitive abilities, including cumulative culture and language. Shared intentionality rests on a fundamentally prosocial disposition that is strikingly absent in chimpanzees, but present in cooperatively breeding primates. Thus, our hypothesis is that while chimpanzees and perhaps all great apes exhibit many of the important cognitive preconditions for uniquely human mental capacities to evolve, they lack the psychological preconditions. In humans, we argue, the two components merged, the cognitive component due to common descent from ape ancestors and the motivational component due to convergent evolution of traits typical of many cooperative breeders.As recently as 6 to 7 million years ago, the hominin lineage split off from the rest of the great ape (hominid) lineage 1 and consequently shares many biological traits and behavioral and cognitive similarities with great apes. 2,3Nevertheless, humans also exhibit remarkable differences from our closest relatives. First, we not only live far longer lives and reproduce at faster rates than do the other great apes, but our offspring take much longer to mature and women cease reproduction well before somatic senescence sets in.4 Second, our ecological niche involves specialization on large, valuable food packages that have to be acquired together, as well as shared, and mandatory reliance on techniques acquired through cumulative culture.5 This niche and our social relationships are based on an un-apelike selflessness, a degree of hypersociality reflected in a concern for others, eagerness to share food and information with others, and cooperation in a wide array of contexts, even with nonrelatives and near-strangers.5-8 Our mode of life facilitates our spread into new habitats, resulting in a ubiquitous geographic distribution.9 Third, with regard to intellectual performance, humans differ from the other great apes, which, as a group, show relatively homogeneous cognitive abilities. [10][11][12]

For over a century, various neuroanatomical measures have been employed as assays of cognitive ability in comparative studies. Nevertheless, it is still unclear whether these measures actually correspond to cognitive ability. A recent meta-analysis of cognitive performance of a broad set of primate species has made it possible to provide a quantitative estimate of general cognitive ability across primates. We find that this estimate is not strongly correlated with neuroanatomical measures that statistically control for a possible effect of body size, such as encephalization quotient or brain size residuals. Instead, absolute brain size measures were the best predictors of primate cognitive ability. Moreover, there was no indication that neocortex-based measures were superior to measures based on the whole brain. The results of previous comparative studies on the evolution of intelligence must be reviewed with this conclusion in mind.

Proactive, that is, unsolicited, prosociality is a key component of our hyper-cooperation, which in turn has enabled the emergence of various uniquely human traits, including complex cognition, morality and cumulative culture and technology. However, the evolutionary foundation of the human prosocial sentiment remains poorly understood, largely because primate data from numerous, often incommensurable testing paradigms do not provide an adequate basis for formal tests of the various functional hypotheses. We therefore present the results of standardized prosociality experiments in 24 groups of 15 primate species, including humans. Extensive allomaternal care is by far the best predictor of interspecific variation in proactive prosociality. Proactive prosocial motivations therefore systematically arise whenever selection favours the evolution of cooperative breeding. Because the human data fit this general primate pattern, the adoption of cooperative breeding by our hominin ancestors also provides the most parsimonious explanation for the origin of human hyper-cooperation.

Human cooperation is unparalleled in the animal world and rests on an altruistic concern for the welfare of genetically unrelated strangers. The evolutionary roots of human altruism, however, remain poorly understood. Recent evidence suggests a discontinuity between humans and other primates because individual chimpanzees do not spontaneously provide food to other group members, indicating a lack of concern for their welfare. Here, we demonstrate that common marmoset monkeys (Callithrix jacchus) do spontaneously provide food to nonreciprocating and genetically unrelated individuals, indicating that other-regarding preferences are not unique to humans and that their evolution did not require advanced cognitive abilities such as theory of mind. Because humans and marmosets are cooperative breeders and the only two primate taxa in which such unsolicited prosociality has been found, we conclude that these prosocial predispositions may emanate from cooperative breeding.cooperative breeding ͉ non-human primates ͉ prosociality ͉ helping ͉ dictator game