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We describe our recent studies of imitation and cultural transmission in chimpanzees and children, which question late twentieth-century characterizations of children as imitators, but chimpanzees as emulators. As emulation entails learning only about the results of others' actions, it has been thought to curtail any capacity to sustain cultures. Recent chimpanzee diffusion experiments have by contrast documented a significant capacity for copying local behavioural traditions. Additionally, in recent ‘ghost’ experiments with no model visible, chimpanzees failed to replicate the object movements on which emulation is supposed to focus. We conclude that chimpanzees rely more on imitation and have greater cultural capacities than previously acknowledged. However, we also find that they selectively apply a range of social learning processes that include emulation. Recent studies demonstrating surprisingly unselective ‘over-imitation’ in children suggest that children's propensity to imitate has been underestimated too. We discuss the implications of these developments for the nature of social learning and culture in the two species. Finally, our new experiments directly address cumulative cultural learning. Initial results demonstrate a relative conservatism and conformity in chimpanzees' learning, contrasting with cumulative cultural learning in young children. This difference may contribute much to the contrast in these species' capacities for cultural evolution.

Network optimality has been described in genes, proteins and human communicative networks. In the latter, optimality leads to the efficient transmission of information with a minimum number of connections. Whilst studies show that differences in centrality exist in animal networks with central individuals having higher fitness, network efficiency has never been studied in animal groups. Here we studied 78 groups of primates (24 species). We found that group size and neocortex ratio were correlated with network efficiency. Centralisation (whether several individuals are central in the group) and modularity (how a group is clustered) had opposing effects on network efficiency, showing that tolerant species have more efficient networks. Such network properties affecting individual fitness could be shaped by natural selection. Our results are in accordance with the social brain and cultural intelligence hypotheses, which suggest that the importance of network efficiency and information flow through social learning relates to cognitive abilities.

Animal ambassador programs are increasingly prevalent in zoos, yet few studies have investigated their impact on animal welfare. We assessed the effects of an ambassador program on the behavior of a colony (N = 15) of zoo-housed African penguins (Spheniscus demersus) and evaluated whether individual characteristics were predictive of participation. Behavioral data were collected for 16 weeks and included 43 “penguin encounters”, during which zoo visitors entered a designated portion of the penguins’ enclosure. When comparing colony behavior following encounters to behavior during a matched control period lacking an encounter, we found no significant difference between affiliative or aggressive behaviors, suggesting that the encounters did not disrupt interactions in the colony. The same was true when comparing behavior preceding the encounter to a matched control period, indicating that any anticipatory period was similarly non-disruptive. Space use during encounters suggested comfort near visitors. We also measured penguin temperament on the shy-bold continuum by recording the birds’ response to novel objects and found that penguins’ temperament, sex, and age were predictive of participation. We concluded that this program had a neutral or positive impact on penguin welfare and considered the findings in relation to aspects of the ambassador program that provided penguins with control over their involvement.

Studying animal cognition in a social setting is associated with practical and statistical challenges. However, conducting cognitive research without disturbing species-typical social groups can increase ecological validity, minimize distress, and improve animal welfare. Here, we review the existing literature on cognitive research run with primates in a social setting in order to determine how widespread such testing is and highlight approaches that may guide future research planning. Using Google Scholar to search the terms \"primate\" \"cognition\" \"experiment\" and \"social group,\" we conducted a systematic literature search covering 16 years (2000-2015 inclusive). We then conducted two supplemental searches within each journal that contained a publication meeting our criteria in the original search, using the terms \"primate\" and \"playback\" in one search and the terms \"primate\" \"cognition\" and \"social group\" in the second. The results were used to assess how frequently nonhuman primate cognition has been studied in a social setting (>3 individuals), to gain perspective on the species and topics that have been studied, and to extract successful approaches for social testing. Our search revealed 248 unique publications in 43 journals encompassing 71 species. The absolute number of publications has increased over years, suggesting viable strategies for studying cognition in social settings. While a wide range of species were studied they were not equally represented, with 19% of the publications reporting data for chimpanzees. Field sites were the most common environment for experiments run in social groups of primates, accounting for more than half of the results. Approaches to mitigating the practical and statistical challenges were identified. This analysis has revealed that the study of primate cognition in a social setting is increasing and taking place across a range of environments. This literature review calls attention to examples that may provide valuable models for researchers wishing to overcome potential practical and statistical challenges to studying cognition in a social setting, ultimately increasing validity and improving the welfare of the primates we study.

Among the growing list of novel tools with which to assess animal welfare is the use of thermal (infrared) imaging. The technology has already been utilized to identify emotional arousal in several nonhuman primate species, though most of these approaches have necessitated the use of relatively controlled settings. Here, we were interested to determine the feasibility of such techniques in a sanctuary setting in which chimpanzees were unrestrained and able to move freely around their enclosures. Furthermore, we sought to evaluate how such thermal images could be paired with corresponding long-term behavioral data and contribute to a multifactorial welfare monitoring system. Over a 6-month period, we simultaneously collected both behavioral and thermographic data on 29 chimpanzees living in four social groups. While we took a thermal image with every behavioral data point, we found that only a small proportion (6.38%) of the thermal images we captured were of sufficient quality to analyze. Most of these usable thermal images (55%) corresponded with a behavioral observation scored as “inactive,” and thus other, less frequent behaviors are not so well represented in our final data set. From our data set, we were able to determine that nasal temperatures were relatively lower when chimpanzees were categorized in active behaviors compared to inactive behaviors, providing some validity measures to our approach. While there are other potential applications for thermal imaging in the behavioral management of chimpanzees, managers should consider the practical limitations of developing long-term welfare monitoring programs that rely on thermographic data.

In contrast to reports of wild primates, studies of captive primates’ flexibility often reveal conservatism: individuals are unable to switch to new and more efficient strategies when task demands change. We propose that such conservatism might be a result of task design and hypothesize that conservatism might be linked to primates’ lack of causal understanding in relation to experimental apparatuses. We investigated if chimpanzees ( Pan troglodytes ) and western lowland gorillas ( Gorilla gorilla gorilla ) would show greater flexibility when presented with a causally-clear task. We presented six chimpanzees and seven gorillas with a clear tube from which they had to remove straws to release a reward. To first evaluate the apes’ causal understanding, we recorded the efficiency with which the apes solved the task (i.e., whether they only removed straws below the reward, ignoring redundant ones above it). To further explore how they solved the task, we also recorded the order in which they removed the straws, which allowed us to determine if habitual action sequences emerged. All apes spontaneously solved the task in their first trial and across repeated trials the majority of their solutions were efficient (median = 90.9%), demonstrating their understanding of the puzzle. There was individual variation in the consistency of straw removal patterns exhibited by the apes, but no ape developed an exclusive habit in the order with which they removed the straws, further indicating their causal understanding of the task. Next, we presented the apes with a new configuration of the same task that required the apes to remove fewer straws to obtain the reward. All apes switched to a more efficient straw removal sequence even though their previously-successful, but now less-efficient, solution remained available. We theorize that because the apes understood the causality of the task, they did not form habits and were not conservative.

The Stroop effect describes interference in cognitive processing due to competing cognitive demands. Presenting emotionally laden stimuli creates similar Stroop-like effects that result from participants’ attention being drawn to distractor stimuli. Here, we adapted the methods of a pictorial Stroop study for use with chimpanzees (N = 6), gorillas (N = 7), and Japanese macaques (N = 6). We tested all subjects via touchscreens following the same protocol. Ten of the 19 subjects passed pre-test training. Subjects who reached criterion were then tested on a standard color-interference Stroop test, which revealed differential accuracy in the primates’ responses across conditions. Next, to test for an emotional Stroop effect, we presented subjects with photographs that were either positively valenced (a preferred food) or negatively valenced (snakes). In the emotional Stroop task, as predicted, the primates were less accurate in trials which presented emotionally laden stimuli as compared to control trials, but there were differences in the apes’ and monkeys’ response patterns. Furthermore, for both Stroop tests, while we found that subjects’ accuracy rates were reduced by test stimuli, in contrast to previous research, we found no difference across trial types in the subjects’ response latencies across conditions.

Nonhuman primates are more likely to learn from the actions of a social model than a non-social “ghost display”, however the mechanism underlying this effect is still unknown. One possibility is that live models are more engaging, drawing increased attention to social stimuli. However, recent research with humans has suggested that live models fundamentally alter memory, not low-level attention. In the current study, we developed a novel eye-tracking paradigm to disentangle the influence of social context on attention and memory in apes. Tested in two conditions, zoo-housed apes (2 gorillas, 5 chimpanzees) were familiarized to videos of a human hand (social condition) and mechanical claw (non-social condition) constructing a three-block tower. During the memory test, subjects viewed side-by-side pictures of the previously-constructed block tower and a novel block tower. In accordance with looking-time paradigms, increased looking time to the novel block tower was used to measure event memory. Apes evidenced memory for the event featuring a social model, though not for the non-social condition. This effect was not dependent on attention differences to the videos. These findings provide the first evidence that, like humans, social stimuli increase nonhuman primates’ event memory, which may aid in information transmission via social learning.

Despite the importance of individual problem solvers for group- and individual-level fitness, the correlates of individual problem-solving success are still an open topic of investigation. In addition to demographic factors, such as age or sex, certain personality dimensions have also been revealed as reliable correlates of problem-solving by animals. Such correlates, however, have been little-studied in chimpanzees. To empirically test the influence of age, sex, estrous state, and different personality factors on chimpanzee problem-solving, we individually tested 36 captive chimpanzees with two novel foraging puzzles. We included both female ( N  = 24) and male ( N  = 12) adult chimpanzees (aged 14–47 years) in our sample. We also controlled for the females’ estrous state—a potential influence on cognitive reasoning—by testing cycling females both when their sexual swelling was maximally tumescent (associated with the luteinizing hormone surge of a female’s estrous cycle) and again when it was detumescent. Although we found no correlation between the chimpanzees’ success with either puzzle and their age or sex, the chimpanzees’ personality ratings did correlate with responses to the novel foraging puzzles. Specifically, male chimpanzees that were rated highly on the factors Methodical, Openness (to experience), and Dominance spent longer interacting with the puzzles. There was also a positive relationship between the latency of females to begin interacting with the two tasks and their rating on the factor Reactivity/Undependability. No other significant correlations were found, but we report tentative evidence for increased problem-solving success by the females when they had detumescent estrous swellings.

Wild chimpanzees frequently make arboreal nests, while wild lowland gorillas typically nest on the ground. We aimed to understand whether zoo-housed apes’ use of elevated spaces for retiring similarly differed between species and across exhibits. Using a pre-planned exhibit switch at Lincoln Park Zoo (Chicago, USA), we compared where (elevated or terrestrial) two groups of apes (Pan troglodytes and Gorilla gorilla gorilla) performed retiring behaviors (inactive, sleeping, and nest-building behaviors). We studied a group of six chimpanzees and a group of four gorillas in two exhibits of similar size and configuration for two three-month periods (between 3 and 5 p.m.) before and after the groups switched exhibits. We predicted that chimpanzees would be more likely to retire in elevated locations compared to gorillas, irrespective of the exhibit. We found a significant effect of exhibit on where the apes retired but no effect of species, such that both species were more likely to retire in elevated locations in one exhibit but not the other. This suggests that the specific characteristics of the exhibits (e.g., number of visual barriers) influenced the expression of the apes’ retiring behaviors. These findings offer further insight in how exhibit design can influence the expression of natural behaviors in these species.