Innovative multi-material tool use in the pant-hoot display of a chimpanzee

‘Pant-hoot displays’ are a species-typical, multi-modal communicative behaviour in chimpanzees in which pant-hoot vocalisations are combined with varied behavioural displays. In both captivity and the wild, individuals commonly incorporate striking or throwing elements of their environment into these displays. In this case study, we present five videos of an unenculturated, captive, adult male chimpanzee combining a large rubber feeding tub with excelsior (wood wool) in a multi-step process, which was then integrated into the subject’s pant-hoot displays as a percussive tool or ‘instrument’. During the construction process, the subject demonstrated an understanding of the relevant properties of these materials, ‘repairing’ the tub to be a more functional drum when necessary. We supplement these videos with a survey of care staff from the study site for additional detail and context. Although care must be taken in generalising data from a single individual, the behaviour reported here hints at three intriguing features of chimpanzee communicative cognition: (1) it suggests a degree of voluntary control over vocal production, (2) it is a so-far unique example of compound tool innovation and use in communicative behaviour and (3) it may represent an example of forward planning in communicative behaviour. Each of these would represent hitherto undocumented dimensions of flexibility in chimpanzee communication, mapping fertile ground for future research

Behavioral conservatism is linked to complexity of behavior in chimpanzees (<i>Pan troglodytes</i>):implications for cognition and cumulative culture

Cumulative culture is rare, if not altogether absent in nonhuman species. At the foundation of cumulative learning is the ability to modify, relinquish, or build upon previous behaviors flexibly to make them more productive or efficient. Within the primate literature, a failure to optimize solutions in this way is often proposed to derive from low-fidelity copying of witnessed behaviors, suboptimal social learning heuristics, or a lack of relevant sociocognitive adaptations. However, humans can also be markedly inflexible in their behaviors, perseverating with, or becoming fixated on, outdated or inappropriate responses. Humans show differential patterns of flexibility as a function of cognitive load, exhibiting difficulties with inhibiting suboptimal behaviors when there are high demands on working memory. We present a series of studies on captive chimpanzees that indicate that behavioral conservatism in apes may be underlain by similar constraints: Chimpanzees showed relatively little conservatism when behavioral optimization involved the inhibition of a well-established but simple solution, or the addition of a simple modification to a well-established but complex solution. In contrast, when behavioral optimization involved the inhibition of a well-established but complex solution, chimpanzees showed evidence of conservatism. We propose that conservatism is linked to behavioral complexity, potentially mediated by cognitive resource availability, and may be an important factor in the evolution of cumulative culture.</p

Chimpanzees prioritise social information over pre-existing behaviours in a group context but not in dyads

Funding: John Templeton Foundation (US) (40128).How animal communities arrive at homogeneous behavioural preferences is a central question for studies of cultural evolution. Here, we investigated whether chimpanzees (Pan troglodytes) would relinquish a pre-existing behaviour to adopt an alternative demonstrated by an overwhelming majority of group mates; in other words, whether chimpanzees behave in a conformist manner. In each of five groups of chimpanzees (N = 37), one individual was trained on one method of opening a two-action puzzle box to obtain food, while the remaining individuals learned the alternative method. Over 5 h of open access to the apparatus in a group context, it was found that 4/5 ‘minority’ individuals explored the majority method and three of these used this new method in the majority of trials. Those that switched did so after observing only a small subset of their group, thereby not matching conventional definitions of conformity. In a further ‘Dyad’ condition, six pairs of chimpanzees were trained on alternative methods and then given access to the task together. Only one of these individuals ever switched method. The number of observations that individuals in the minority and Dyad individuals made of their untrained method was not found to influence whether or not they themselves switched to use it. In a final ‘Asocial’ condition, individuals (N = 10) did not receive social information and did not deviate from their first-learned method. We argue that these results demonstrate an important influence of social context upon prioritisation of social information over pre-existing methods, which can result in group homogeneity of behaviour.Publisher PDFPeer reviewe

Chimpanzees’ Socially Maintained Food Preferences Indicate both Conservatism and Conformity

Chimpanzees remain fixed on a single strategy, even if a novel, more efficient, strategy is introduced. Previous studies reporting such findings have incorporated paradigms in which chimpanzees learn one behavioural method and then are shown a new one that the chimpanzees invariably do not adopt. This study provides the first evidence that chimpanzees show such conservatism even when the new method employs the identical required behaviour as the first, but for a different reward. Groups of chimpanzees could choose to exchange one of two inedible tokens; one was rewarded with a highly preferred food (grape) and the other with a less preferred food (carrot). Individuals first observed a model chimpanzee from their social group trained to choose one of the two types of tokens. In one group, this token earned a carrot, while in the other, control, group the token earned a grape. In both groups, chimpanzees conformed to the trained model’s choice. This was especially striking for those gaining the pieces of carrot; the less favoured reward. This resulted in a population-level trend of food choices, even when counter to their original, individual, preferences. Moreover, the chimpanzees’ food preferences did not change over time, demonstrating that these results were not due to a simple shift in preferences. We discuss social factors apparent in the interactions and suggest that, despite seeming to be inefficient, in chimpanzees, conformity may benefit them, possibly by assisting with the maintenance of group relations

The Ontogeny of Social Comparisons by Rhesus Macaques (Macaca mulatta)

This longitudinal study investigated the development of social contrast-negative responses to inequitable rewards-in rhesus macaques (Macaca mulatta). Although responses to inequity by humans appear universal, this is something that develops with age. Infants first recognize inequity when around 18 months old and respond to it only when they are around 3 years old. To date, however, there have been no studies of the ontogeny of the inequity response in any species other than humans. To address this, we used an exchange paradigm, in which 10 pairs of rhesus monkeys had to exchange inedible tokens with the experimenter to get food rewards that differed in quality depending on the condition. All subjects were tested first when they were an average of 17 months old and a subset, of four pairs, was tested again a year later. Subjects responded negatively to contrast-recognizing a disparity in expected, as compared to, received rewards-based on both social and individual comparisons at the older age, but not at the younger age. Similar to humans, rhesus showed a developmental trajectory to social comparison, providing the first evidence for the ontogeny of this response in a non-human species

When Given the Opportunity, Chimpanzees Maximize Personal Gain Rather than “Level the Playing Field”

We provided chimpanzees (Pan troglodytes) with the ability to improve the quality of food rewards they received in a dyadic test of inequity.We were interested to see if this provision influenced their responses and, if so, whether it was mediated by a social partner’s outcomes. We tested eight dyads using an exchange paradigm in which, depending on the condition, the chimpanzees were rewarded with either high-value (a grape) or low-value (a piece of celery) food rewards for each completed exchange. We included four conditions. In the first, “Different” condition, the subject received different, less-preferred, rewards than their partner for each exchange made (a test of inequity). In the “Unavailable” condition, high-value rewards were shown, but not given, to both chimpanzees prior to each exchange and the chimpanzees were rewarded equally with low-value rewards (a test of individual contrast). The final two conditions created equity. In these High-value and Low-value “Same” conditions both chimpanzees received the same food rewards for each exchange.Within each condition, the chimpanzees first completed ten trials in the Baseline Phase, in which the experimenter determined the rewards they received, and then ten trials in the Test Phase. In the Test Phase, the chimpanzees could exchange tokens through the aperture of a small wooden picture frame hung on their cage mesh in order to receive the high-value reward. Thus, in the Test Phase, the chimpanzees were provided with an opportunity to improve the quality of the rewards they received, either absolutely or relative to what their partner received. The chimpanzees responded in a targeted manner; in the Test Phase they attempted to maximize their returns in all conditions in which they had received low-value rewards during the Baseline Phase. Thus, the chimpanzees were apparently motivated to increase their reward regardless of their partners’, but they only used the mechanism provided when it afforded the opportunity for them to increase their rewards.We also found evidence that the chimpanzees’ responses were enhanced by social facilitation. Specifically, the chimpanzees were more likely to exchange their tokens through the frame when their test partner also did so, even in circumstances in which their reward value could not be improved. Our paradigm provided the chimpanzees with the possibility to improve the quality of rewards they received in the Test Phase. We found that refusals – to exchange tokens or to eat rewards – decreased significantly in the Test Phase compared to the Baseline Phase, where no such opportunity for improvement of outcomes existed. Thus, the chimpanzees participated more when they could improve the rewards they received

Mechanisms Underlying the Response to Inequitable Outcomes in Chimpanzees, Pan Troglodytes

Several species of non-human primates respond negatively to inequitable outcomes, a trait shared with humans. Despite previous research, questions regarding the response to inequity remain. In this study, we replicated the methodology from previous studies to address four questions related to inequity. First, we explored the impact of basic social factors. Second, we addressed whether negative responses to inequity require a task, or exist when rewards are given for ‘free’. Third, we addressed whether differences in the experimental procedure or the level of effort required to obtain a reward affected responses. Finally, we explored the interaction between ‘individual’ expectations (based on one’s own previous experience) and ‘social’ expectations (based on the partner’s experience). These questions were investigated in 16 socially-housed adult chimpanzees using eight conditions that varied across the dimensions of reward, effort, and procedure. Subjects did respond to inequity, but only in the context of a task. Differences in procedure and level of effort required did not cause individuals to change their behavior. Males were more sensitive to social than to individual expectation, while females were more sensitive to individual expectation. Finally, subjects also increased refusals when receiving a better reward than their partner, which has not been seen previously. These results indicate that chimpanzees are more sensitive to reward inequity than procedures, and that there is interaction between social and individual expectations that depends upon social factors

Foundations of cumulative culture in apes: improved foraging efficiency through relinquishing and combining witnessed behaviours in chimpanzees (Pan troglodytes)

This research was funded by the John Templeton Foundation (Grant ID: 40128, to K. Laland and A. Whiten).A vital prerequisite for cumulative culture, a phenomenon often asserted to be unique to humans, is the ability to modify behaviour and flexibly switch to more productive or efficient alternatives. Here, we first established an inefficient solution to a foraging task in five captive chimpanzee groups (N = 19). Three groups subsequently witnessed a conspecific using an alternative, more efficient, solution. When participants could successfully forage with their established behaviours, most individuals did not switch to this more efficient technique; however, when their foraging method became substantially less efficient, nine chimpanzees with socially-acquired information (four of whom witnessed additional human demonstrations) relinquished their old behaviour in favour of the more efficient one. Only a single chimpanzee in control groups, who had not witnessed a knowledgeable model, discovered this. Individuals who switched were later able to combine components of their two learned techniques to produce a more efficient solution than their extensively used, original foraging method. These results suggest that, although chimpanzees show a considerable degree of conservatism, they also have an ability to combine independent behaviours to produce efficient compound action sequences; one of the foundational abilities (or candidate mechanisms) for human cumulative culture.Publisher PDFPeer reviewe

Evolution and the Expression of Biases: Situational Value Changes the Endowment Effect in Chimpanzees

Cognitive and behavioral biases, which are widespread among humans, have recently been demonstrated in other primates, suggesting a common origin. Here we examine whether the expression of one shared bias, the endowment effect, varies as a function of context. We tested whether objects lacking inherent value elicited a stronger endowment effect (or preference for keeping the object) in a context in which the objects had immediate instrumental value for obtaining valuable resources (food). Chimpanzee subjects had opportunities to trade tools when food was not present, visible but unobtainable, and obtainable using the tools. We found that the endowment effect for these tools existed only when they were immediately useful, showing that the effect varies as a function of context-specific utility. Such context-specific variation suggests that the variation seen in some human biases may trace predictably to behaviors that evolved to maximize gains in specific circumstances

Evolution and the Expression of Biases: Situational Value Changes the Endowment Effect

Cognitive and behavioral biases, which are widespread among humans, have recently been demonstrated in other primates, suggesting a common origin. Here we examine whether the expression of one shared bias, the endowment effect, varies as a function of context. We tested whether objects lacking inherent value elicited a stronger endowment effect (or preference for keeping the object) in a context in which the objects had immediate instrumental value for obtaining valuable resources (food). Chimpanzee subjects had opportunities to trade tools when food was not present, visible but unobtainable, and obtainable using the tools. We found that the endowment effect for these tools existed only when they were useful, showing that the effect varies as a function of context-specific utility. Such context-specific variation suggests that the variation seen in some human biases may trace predictably to behaviors that evolved to maximize gains in specific circumstances