Bonobos and chimpanzees remember familiar conspecifics for decades.

Recognition and memory of familiar conspecifics provides the foundation for complex sociality and is vital to navigating an unpredictable social world [Tibbetts and Dale, Trends Ecol. Evol. 22, 529-537 (2007)]. Human social memory incorporates content about interactions and relationships and can last for decades [Sherry and Schacter, Psychol. Rev. 94, 439-454 (1987)]. Long-term social memory likely played a key role throughout human evolution, as our ancestors increasingly built relationships that operated across distant space and time [Malone et al., Int. J. Primatol. 33, 1251-1277 (2012)]. Although individual recognition is widespread among animals and sometimes lasts for years, little is known about social memory in nonhuman apes and the shared evolutionary foundations of human social memory. In a preferential-looking eye-tracking task, we presented chimpanzees and bonobos (N = 26) with side-by-side images of a previous groupmate and a conspecific stranger of the same sex. Apes' attention was biased toward former groupmates, indicating long-term memory for past social partners. The strength of biases toward former groupmates was not impacted by the duration apart, and our results suggest that recognition may persist for at least 26 y beyond separation. We also found significant but weak evidence that, like humans, apes may remember the quality or content of these past relationships: apes' looking biases were stronger for individuals with whom they had more positive histories of social interaction. Long-lasting social memory likely provided key foundations for the evolution of human culture and sociality as they extended across time, space, and group boundaries.

Chimpanzees (Pan troglodytes) recognize that their guesses could be wrong and can pass a two-cup disjunctive syllogism task.

When chimpanzees search for hidden food, do they realize that their guesses may not be correct? We applied a post-decision wagering paradigm to a simple two-cup search task, varying whether we gave participants visual access to the baiting and then asking after they had chosen one of the cups whether they would prefer a smaller but certain reward instead of their original choice (experiment 1). Results showed that chimpanzees were more likely to accept the smaller reward in occluded than visible conditions. Experiment 2 found the same effect when we blocked visual access but manipulated the number of hiding locations for the food piece, showing that the effect is not owing to representation type. Experiments 3 and 4 showed that when given information about the contents of the unchosen cup, chimpanzees were able to flexibly update their choice behaviour accordingly. These results suggest that language is not a pre-requisite to solving the disjunctive syllogism and provides a valuable contribution to the debate on logical reasoning in non-human animals.

Orcas remember what to copy: a deferred and interference-resistant imitation study.

Response facilitation has often been portrayed as a "low level" category of social learning, because the demonstrator's action, which is already in the observer's repertoire, automatically triggers that same action, rather than induces the learning of a new action. One way to rule out response facilitation consists of introducing a delay between the demonstrator's behavior and the observer's response to let their possible effects wear off. However, this may not rule out "delayed response facilitation" in which the subject could be continuously "mentally rehearsing" the demonstrated actions during the waiting period. We used a do-as-the-other-did paradigm in two orcas to study whether they displayed cognitive control regarding their production of familiar actions by (1) introducing a delay ranging from 60 to 150 s between observing and producing the actions and (2) interspersing distractor (non-target) actions performed by the demonstrator and by the subjects during the delay period. These two manipulations were aimed at preventing the mental rehearsal of the observed actions during the delay period. Both orcas copied the model's target actions on command after various delay periods, and crucially, despite the presence of distractor actions. These findings suggest that orcas are capable of selectively retrieving a representation of an observed action to generate a delayed matching response. Moreover, these results lend further support to the proposal that the subjects' performance relied not only on a mental representation of the specific actions that were requested to copy, but also flexibly on the abstract and domain general rule requested by the specific "copy command". Our findings strengthen the view that orcas and other cetaceans are capable of flexible and controlled social learning.

Chimpanzees prepare for alternative possible outcomes.

When facing uncertainty, humans often build mental models of alternative outcomes. Considering diverging scenarios allows agents to respond adaptively to different actual worlds by developing contingency plans (covering one's bases). In a pre-registered experiment, we tested whether chimpanzees (Pan troglodytes) prepare for two mutually exclusive possibilities. Chimpanzees could access two pieces of food, but only if they successfully protected them from a human competitor. In one condition, chimpanzees could be certain about which piece of food the human experimenter would attempt to steal. In a second condition, either one of the food rewards was a potential target of the competitor. We found that chimpanzees were significantly more likely to protect both pieces of food in the second relative to the first condition, raising the possibility that chimpanzees represent and prepare effectively for different possible worlds.

Do chimpanzees reason logically?

Psychologists disagree about the development of logical concepts such as or and not. While some theorists argue that infants reason logically, others maintain that logical inference is contingent on linguistic abilities and emerges around age 4. In this Registered Report, we conducted five experiments on logical reasoning in chimpanzees. Subjects (N = 16; 10 females; M = 24 years) participated in the same setup that has been administered to children: the two-, three-, and four-cup-task. Chimpanzees performed above chance in the two-cup-, but not in the three-cup-task. Furthermore, chimpanzees selected the logically correct option more often in the test than the control condition of the four-cup-task. We discuss possible interpretations of these findings and conclude that our results are most consistent with non-deductive accounts.

On the murky dissociation between expression and communication.

The authors present an ambitious attempt to outline the gradual evolution of the cognitive foundations of ostensive communication. We focus on three problematic aspects of the distinction between expression and communication: ambiguity in the distinction's central principle of "complementary mechanisms," inconsistencies in the application of the distinction across taxa, and the dismissal of mentalizing in nonhuman primates.

Human adults prefer to cooperate even when it is costly.

Joint actions are cooperative activities where humans coordinate their actions to achieve individual and shared goals. While the motivation to engage in joint action is clear when a goal cannot be achieved by individuals alone, we asked whether humans are motivated to act together even when acting together is not necessary and implies incurring additional costs compared to individual goal achievement. Using a utility-based empirical approach, we investigated the extent of humans' preference for joint action over individual action, when the instrumental costs of performing joint actions outweigh the benefits. The results of five experiments showed that human adults have a stable preference for joint action, even if individual action is more effective to achieve a certain goal. We propose that such preferences can be understood as ascribing additional reward value to performing actions together.

Great apes and human children rationally monitor their decisions.

Several species can detect when they are uncertain about what decision to make-revealed by opting out of the choice, or by seeking more information before deciding. However, we do not know whether any nonhuman animals recognize when they need more information to make a decision because new evidence contradicts an already-formed belief. Here, we explore this ability in great apes and human children. First, we show that after great apes saw new evidence contradicting their belief about which of two rewards was greater, they stopped to recheck the evidence for their belief before deciding. This indicates the ability to keep track of the reasons for their decisions, or 'rational monitoring' of the decision-making process. Children did the same at 5 years of age, but not at 3 years. In a second study, participants formed a belief about a reward's location, but then a social partner contradicted them, by picking the opposite location. This time even 3-year-old children rechecked the evidence, while apes ignored the disagreement. While apes were sensitive only to the conflict in physical evidence, the youngest children were more sensitive to peer disagreement than conflicting physical evidence.

Understanding Language Evolution: Beyond Pan-Centrism.

Language does not fossilize but this does not mean that the language's evolutionary timeline is lost forever. Great apes provide a window back in time on our last prelinguistic ancestor's communication and cognition. Phylogeny and cladistics implicitly conjure Pan (chimpanzees, bonobos) as a superior (often the only) model for language evolution compared with earlier diverging lineages, Gorilla and Pongo (orangutans). Here, in reviewing the literature, it is shown that Pan do not surpass other great apes along genetic, cognitive, ecologic, or vocal traits that are putatively paramount for language onset and evolution. Instead, revived herein is the idea that only by abandoning single-species models and learning about the variation among great apes, there might be a chance to retrieve lost fragments of the evolutionary timeline of language.

Exploring the development of attentional set shifting in young children with a novel Intradimensional/Extradimensional shift task.

Attentional set shifting is a core part of cognition, allowing quick and flexible adaption to new demands. The study of its development during early childhood has been hampered by a shortage of measures not requiring language. This article argues for a revival of the Intradimensional/Extradimensional (ID/ED) shift task by presenting a new nonverbal version of the task (Shifting Tray task). Children (N = 95 3- to 5-year-olds; 49 girls; predominantly European White) were presented with pairs of trays, each filled with a substrate and an upside-down cup on top, and were asked to find stickers. In the pre-switch phase, children learned (through trial and error) which dimension (substrate or cup) was predictive of the rewards. In the post-switch phase, all stimuli were exchanged. For children in the intradimensional shift condition, the dimension predictive of the sticker was the same as the one predictive in the pre-switch phase. For children in the extradimensional shift condition, the previously irrelevant dimension was now relevant. Results showed that most 3-year-olds were able to switch, and older children did not outperform younger children. The easy and flexible nature of the task allows researchers to investigate the impact of labels and instructions and to use it in cross-cultural and comparative research.

Primate cognition in zoos: Reviewing the impact of zoo-based research over 15 years.

Primate cognition research is reliant on access to members of the study sp ecies and logistical infrastructures to conduct observations and experiments. Historically founded in research centers and private collections, and spreading to modern zoos, sanctuaries, and the field, primate cognition has been investigated in diverse settings, each with benefits and challenges. In our systematic review of 12 primatology, animal behavior, and animal cognition journals over the last 15 years, we turn a spotlight on zoos to quantify their current impact on the field and to highlight their potential as robust contributors to future work. To put zoo-based research in context, we compare zoos to three other site types: university-owned or independent research centers, sanctuaries, and field sites. We assess the contributions of zoos across several critical considerations in primate cognition research, including number of investigations, species diversity, sample size, research topic diversity, and methodology. We identified 1119 publications reporting studies of primate cognition, almost 25% of which report research conducted in zoos. Across publications, zoo-based research has greater species diversity than research centers and covers a diverse range of research topics. Although our review is merely a snapshot of primate cognition research, our findings suggest that zoos may present advantages to researchers regarding species diversity, and lack some of the methodological constraints of field sites, allowing greater ease of access to a diverse range of subjects for cognition investigations. We suggest that zoos have great potential as key contributors for future investigations in primate cognition. Finally, we shed light on the symbiotic relationship that can emerge between researchers and zoos, forming partnerships that bring unique advantages to both parties.

Great apes use self-experience to anticipate an agent's action in a false-belief test.

Human social life depends on theory of mind, the ability to attribute mental states to oneself and others. A signature of theory of mind, false belief understanding, requires representing others' views of the world, even when they conflict with one's own. After decades of research, it remains controversial whether any nonhuman species possess a theory of mind. One challenge to positive evidence of animal theory of mind, the behavior-rule account, holds that animals solve such tasks by responding to others' behavioral cues rather than their mental states. We distinguish these hypotheses by implementing a version of the "goggles" test, which asks whether, in the absence of any additional behavioral cues, animals can use their own self-experience of a novel barrier being translucent or opaque to determine whether another agent can see through the same barrier. We incorporated this paradigm into an established anticipatory-looking false-belief test for great apes. In a between-subjects design, apes experienced a novel barrier as either translucent or opaque, although both looked identical from afar. While being eye tracked, all apes then watched a video in which an actor saw an object hidden under 1 of 2 identical boxes. The actor then scuttled behind the novel barrier, at which point the object was relocated and then removed. Only apes who experienced the barrier as opaque visually anticipated that the actor would mistakenly search for the object in its previous location. Great apes, therefore, appeared to attribute differential visual access based specifically on their own past perceptual experience to anticipate an agent's actions in a false-belief test.

Evolutionary foundations of knowledge and belief attribution in nonhuman primates.

Recent findings from anticipatory-looking false-belief tests have shown that nonhuman great apes and macaques anticipate that an agent will go to the location where the agent falsely believed an object to be. Phillips et al.'s claim that nonhuman primates attribute knowledge but not belief should thus be reconsidered. We propose that both knowledge and belief attributions are evolutionary old.

Learning Novel Skills From Iconic Gestures: A Developmental and Evolutionary Perspective.

Cumulative cultural learning has been argued to rely on high-fidelity copying of other individuals' actions. Iconic gestures of actions have no physical effect on objects in the world but merely represent actions that would have an effect. Learning from iconic gestures thus requires paying close attention to the teacher's precise bodily movements-a prerequisite for high-fidelity copying. In three studies, we investigated whether 2- and 3-year-old children (N = 122) and great apes (N = 36) learn novel skills from iconic gestures. When faced with a novel apparatus, participants watched an experimenter perform either an iconic gesture depicting the action necessary to open the apparatus or a gesture depicting a different action. Children, but not great apes, profited from iconic gestures, with older children doing so to a larger extent. These results suggest that high-fidelity copying abilities are firmly in place in humans by at least 3 years of age.

Age influences domestic dog cognitive performance independent of average breed lifespan.

Across mammals, increased body size is positively associated with lifespan. However, within species, this relationship is inverted. This is well illustrated in dogs (Canis familiaris), where larger dogs exhibit accelerated life trajectories: growing faster and dying younger than smaller dogs. Similarly, some age-associated traits (e.g., growth rate and physiological pace of aging) exhibit accelerated trajectories in larger breeds. Yet, it is unknown whether cognitive performance also demonstrates an accelerated life course trajectory in larger dogs. Here, we measured cognitive development and aging in a cross-sectional study of over 4000 dogs from 66 breeds using nine memory and decision-making tasks performed by citizen scientists as part of the Dognition project. Specifically, we tested whether cognitive traits follow a compressed (accelerated) trajectory in larger dogs, or the same trajectory for all breeds, which would result in limited cognitive decline in larger breeds. We found that all breeds, regardless of size or lifespan, tended to follow the same quadratic trajectory of cognitive aging-with a period of cognitive development in early life and decline in later life. Taken together, our results suggest that cognitive performance follows similar age-related trajectories across dog breeds, despite remarkable variation in developmental rates and lifespan.

Do 7-year-old children understand social leverage?

Individuals with an advantageous position during a negotiation possess leverage over their partners. Several studies with adults have investigated how leverage can influence the coordination strategies of individuals when conflicts of interest arise. In this study, we explored how pairs of 7-year-old children solved a coordination game (based on the Snowdrift scenario) when one child had leverage over the other child. We presented a social dilemma in the form of an unequal reward distribution on a rotating tray. The rotating tray could be accessed by both children. The child who waited longer to act received the best outcome, but if both children waited too long, they would lose the rewards. In addition, one child could forgo the access to the rotating tray for an alternative option-the leverage. Although children rarely used their leverage strategically, children with access to the alternative were less likely to play the social dilemma, especially when their leverage was larger. Furthermore, children waited longer to act as the leverage decreased. Finally, children almost never failed to coordinate. The results hint to a trade-off between maximizing benefits while maintaining long-term collaboration in complex scenarios where strategies such as turn taking are hard to implement.

Chimpanzees flexibly update working memory contents and show susceptibility to distraction in the self-ordered search task.

Working memory (WM) is a core executive function that allows individuals to hold, process and manipulate information. WM capacity has been repeatedly nominated as a key factor in human cognitive evolution; nevertheless, little is known about the WM abilities of our closest primate relatives. In this study, we examined signatures of WM ability in chimpanzees (Pan troglodytes). Standard WM tasks for humans (Homo sapiens) often require participants to continuously update their WM. In Experiment 1, we implemented this updating requirement in a foraging situation: zoo-housed chimpanzees (n = 13) searched for food in an array of containers. To avoid redundant searches, they needed to continuously update which containers they had already visited (similar to WM paradigms for human children) with 15 s retention intervals in between each choice. We examined chimpanzees' WM capacity and to what extent they used spatial cues and object features to memorize their previous choices. In Experiment 2, we investigated how susceptible their WM was to attentional interference, an important signature, setting WM in humans apart from long-term memory. We found large individual differences with some individuals remembering at least their last four choices. Chimpanzees used a combination of spatial cues and object features to remember which boxes they had chosen already. Moreover, their performance decreased specifically when competing memory information was introduced. Finally, we found that individual differences in task performance were highly reliable over time. Together, these findings show remarkable similarities between human and chimpanzee WM abilities despite evolutionary and life-history differences.

Thirty years of great ape gestures.

We and our colleagues have been doing studies of great ape gestural communication for more than 30 years. Here we attempt to spell out what we have learned. Some aspects of the process have been reliably established by multiple researchers, for example, its intentional structure and its sensitivity to the attentional state of the recipient. Other aspects are more controversial. We argue here that it is a mistake to assimilate great ape gestures to the species-typical displays of other mammals by claiming that they are fixed action patterns, as there are many differences, including the use of attention-getters. It is also a mistake, we argue, to assimilate great ape gestures to human gestures by claiming that they are used referentially and declaratively in a human-like manner, as apes' "pointing" gesture has many limitations and they do not gesture iconically. Great ape gestures constitute a unique form of primate communication with their own unique qualities.

Innovative problem solving in great apes: the role of visual feedback in the floating peanut task.

Nonhuman great apes show remarkable behavioural flexibility. Some individuals are even able to use water as a tool: They spit water into a vertical tube to make a peanut float upwards until it comes into reach (floating peanut task; FPT). In the current study, we used the FPT to investigate how visual feedback, an end-state demonstration and a social demonstration affect task performance in nonhuman great apes in three experiments. Our results indicate that apes who had acquired the solution with a clear tube maintained it with an opaque one. However, apes starting with an opaque tube failed to solve the task. Additionally, facing the peanut floating on a water-filled tube (i.e., an end-state demonstration) promoted success independent on the availability of visual feedback. Moreover, experiencing how water was poured into the tube either by a human demonstrator or by a water tap that had been opened either by the ape or a human did not seem to be of further assistance. First, this study suggests that great apes require visual feedback for solving the FPT, which is no longer required after the initial acquisition. Second, some subjects benefit from encountering the end-state, a finding corroborating previous studies.

Absolute brain size predicts dog breed differences in executive function.

Large-scale phylogenetic studies of animal cognition have revealed robust links between absolute brain volume and species differences in executive function. However, past comparative samples have been composed largely of primates, which are characterized by evolutionarily derived neural scaling rules. Therefore, it is currently unknown whether positive associations between brain volume and executive function reflect a broad-scale evolutionary phenomenon, or alternatively, a unique consequence of primate brain evolution. Domestic dogs provide a powerful opportunity for investigating this question due to their close genetic relatedness, but vast intraspecific variation. Using citizen science data on more than 7000 purebred dogs from 74 breeds, and controlling for genetic relatedness between breeds, we identify strong relationships between estimated absolute brain weight and breed differences in cognition. Specifically, larger-brained breeds performed significantly better on measures of short-term memory and self-control. However, the relationships between estimated brain weight and other cognitive measures varied widely, supporting domain-specific accounts of cognitive evolution. Our results suggest that evolutionary increases in brain size are positively associated with taxonomic differences in executive function, even in the absence of primate-like neuroanatomy. These findings also suggest that variation between dog breeds may present a powerful model for investigating correlated changes in neuroanatomy and cognition among closely related taxa.