Idiosyncratic gesture use in a mother-infant dyad in chimpanzees (Pan troglodytes schweinfurthii) in the wild.

One promising method to tackle the question, "In which modality did language evolve?" is by studying the ontogenetic trajectory of signals in human's closest living relatives, including chimpanzees (Pan troglodytes). Concerning gestures, current debates centre on four different hypotheses: "phylogenetic ritualization", "social transmission through imitation", "ontogenetic ritualization", and "social negotiation". These differ in their predictions regarding idiosyncratic gestures, making such occurrences a crucial area of investigation. Here, we describe a novel and potential idiosyncratic behaviour - 'hand-on-eye' - which was initially observed in one mother-infant dyad in a community of chimpanzees living in the wild. We systematically investigated the form, sequential organisation, intentionality, usage, function, and distribution of the behaviour over a five-year period. The results showed that 'hand-on-eye' was nearly exclusively deployed in a single mother-infant dyad, was accompanied by hallmarks of intentionality, and served to initiate or resume joint dorsal travel. Although the behaviour was observed once in each of three other mother-infant dyads, these lacked the same frequency and hallmarks of intentionality. 'Hand-on-eye' thus qualifies as an idiosyncratic gesture. The proposed developmental pathway gives support to both the "ontogenetic ritualization" and "social negotiation" hypotheses. It also stresses the crucial need for longitudinal approaches to tackle developmental processes that are triggered by unique circumstances and unfold over relatively long time windows.

Camera traps unveil demography, social structure, and home range of six unhabituated Western chimpanzee groups in the Moyen Bafing National Park, Guinea.

Precise estimates of population dynamics and social grouping patterns are required for effective conservation of wild animal populations. It is difficult to obtain such information on non-human great apes as they have slow reproductive rates. To gain a better understanding of demography in these populations, previous research has typically involved habituation\, a process that requires years. Here, we collected data continuously over year-long periods to monitor an unhabituated population of critically endangered Western chimpanzees (Pan troglodytes verus) in the Moyen Bafing National Park, Guinea. We used two arrays of 100 camera traps that were placed opportunistically in two distinct 100 km2 sites, named Bakoun and Koukoutamba. We identified 227 individuals in Bakoun and 207 in Koukoutamba through their unique facial features. Our camera trap data make clear that these individuals belong to six and seven closed groups, respectively. Six of those groups were near-completely sampled with an average minimum size of 46.8 individuals (range: 37-58), and a mean adult sex ratio of 1.32 (range: 0.93-2.10). We described the demographic composition of these groups and use Bayesian social network analysis to understand population structure. The network analyses suggested that the social bonds within the two populations were structured by sex homophily, with male chimpanzees being more or equally likely to be observed together than other adult associations. Through estimation of minimum convex polygons, we described the minimum home range for those groups. Compared to other chimpanzee groups living in a similar environment (mosaic savanna-forest), the Moyen Bafing region seems to host a high-density of chimpanzees with small home ranges for their group size. Our research highlights the potential of camera traps for studying the demographic composition of chimpanzee populations with high resolution and obtaining crucial information on several groups in a time-efficient and cost-effective way.

Assessing the motivational value of infant video clips on chimpanzees through discrimination learning task.

The motivational value of visual infant stimuli in humans is considered to encourage parental behavior. To explore the evolutionary roots of this preference for infants, we examined the reward value of conspecific infant videos compared to adult ones in nine chimpanzees. We employed a novel approach, a simultaneous discrimination task with differential sensory reinforcement. In Experiments 1 and 2, we tested if watching conspecific infant videos is more rewarding than watching adult ones. Participants were required to discriminate between two visual stimuli by a touch panel task. In video reward trials, a video clip featuring a chimpanzee infant followed a correct choice, while one featuring an adult followed an incorrect choice. However, the percentage of correct choices did not significantly differ from chance except in one chimpanzee, indicating that chimpanzees did not exhibit a preference for watching infant videos over those of adult. In Experiment 3, we tested if chimpanzees prefer conspecific videos over a blank screen; however, we did not find evidence either at a group level. These results suggest that the incentive salience of infant stimuli may not be universally compelling across species. Additionally, we discuss the limitations of the task using sensory reinforcement.

Many morphs: Parsing gesture signals from the noise.

Parsing signals from noise is a general problem for signallers and recipients, and for researchers studying communicative systems. Substantial efforts have been invested in comparing how other species encode information and meaning, and how signalling is structured. However, research depends on identifying and discriminating signals that represent meaningful units of analysis. Early approaches to defining signal repertoires applied top-down approaches, classifying cases into predefined signal types. Recently, more labour-intensive methods have taken a bottom-up approach describing detailed features of each signal and clustering cases based on patterns of similarity in multi-dimensional feature-space that were previously undetectable. Nevertheless, it remains essential to assess whether the resulting repertoires are composed of relevant units from the perspective of the species using them, and redefining repertoires when additional data become available. In this paper we provide a framework that takes data from the largest set of wild chimpanzee (Pan troglodytes) gestures currently available, splitting gesture types at a fine scale based on modifying features of gesture expression using latent class analysis (a model-based cluster detection algorithm for categorical variables), and then determining whether this splitting process reduces uncertainty about the goal or community of the gesture. Our method allows different features of interest to be incorporated into the splitting process, providing substantial future flexibility across, for example, species, populations, and levels of signal granularity. Doing so, we provide a powerful tool allowing researchers interested in gestural communication to establish repertoires of relevant units for subsequent analyses within and between systems of communication.

Apes reciprocate food positively and negatively.

Reciprocal food exchange is widespread in human societies but not among great apes, who may view food mainly as a target for competition. Understanding the similarities and differences between great apes' and humans' willingness to exchange food is important for our models regarding the origins of uniquely human forms of cooperation. Here, we demonstrate in-kind food exchanges in experimental settings with great apes for the first time. The initial sample consisted of 13 chimpanzees and 5 bonobos in the control phases, and the test phases included 10 chimpanzees and 2 bonobos, compared with a sample of 48 human children aged 4 years. First, we replicated prior findings showing no spontaneous food exchanges in great apes. Second, we discovered that when apes believe that conspecifics have 'intentionally' transferred food to them, positive reciprocal food exchanges (food-for-food) are not only possible but reach the same levels as in young children (approx. 75-80%). Third, we found that great apes engage in negative reciprocal food exchanges (no-food for no-food) but to a lower extent than children. This provides evidence for reciprocal food exchange in great apes in experimental settings and suggests that while a potential mechanism of fostering cooperation (via positive reciprocal exchanges) may be shared across species, a stabilizing mechanism (via negative reciprocity) is not.

Chimpanzee and Human Risk Preferences Show Key Similarities.

Risk preference impacts how people make key life decisions related to health, wealth, and well-being. Systematic variations in risk-taking behavior can be the result of differences in fitness expectations, as predicted by life-history theory. Yet the evolutionary roots of human risk-taking behavior remain poorly understood. Here, we studied risk preferences of chimpanzees (86 Pan troglodytes; 47 females; age = 2-40 years) using a multimethod approach that combined observer ratings with behavioral choice experiments. We found that chimpanzees' willingness to take risks shared structural similarities with that of humans. First, chimpanzees' risk preference manifested as a traitlike preference that was consistent across domains and measurements. Second, chimpanzees were ambiguity averse. Third, males were more risk prone than females. Fourth, the appetite for risk showed an inverted-U-shaped relation to age and peaked in young adulthood. Our findings suggest that key dimensions of risk preference appear to emerge independently of the influence of human cultural evolution.

Do chimpanzees see a face on Mars? A search for face pareidolia in chimpanzees.

We sometimes perceive meaningful patterns or images in random arrangements of colors and shapes. This phenomenon is called pareidolia and has recently been studied intensively, especially face pareidolia. In contrast, there are few comparative-cognitive studies on face pareidolia with nonhuman primates. This study explored behavioral evidence for face pareidolia in chimpanzees using visual search and matching tasks. Faces are processed in a configural manner, and their perception and recognition are hampered by inversion and misalignment of top and bottom parts. We investigated whether the same effect occurs in a visual search for face-like objects. The results showed an effect of misalignment. On the other hand, consistent results were not obtained with the photographs of fruits. When only the top or bottom half of the face-like object was presented, chimpanzees showed better performance for the top-half condition, suggesting the importance of the eye area in face pareidolia. In the positive-control experiments, chimpanzees received the same experiment using human faces and human participants with face-like objects and fruits. As a result, chimpanzees showed an inefficient search for inverted and misaligned faces and humans for manipulated face-like objects. Finally, to examine the role of face awareness, we tested matching a human face to a face-like object in chimpanzees but obtained no substantial evidence that they saw the face-like object as a "face." Based on these results, we discussed the extents and limits of face pareidolia in chimpanzees.

Object use in communication of semi-wild chimpanzees.

Object interactions play an important role in human communication but the extent to which nonhuman primates incorporate objects in their social interactions remains unknown. To better understand the evolution of object use, this study explored how objects are used in social interactions in semi-wild chimpanzees (Pan troglodytes). We used an observational approach focusing on naturally occurring object actions where we examined their use and tested whether the production of object actions was influenced by the recipients' visual attention as well as by colony membership. The results show that chimpanzees adjusted both the type of object used, and the modality of object actions to match the visual attention of the recipient, as well as colony differences in the use of targeted object actions. These results provide empirical evidence highlighting that chimpanzees use objects in diverse ways to communicate with conspecifics and that their use may be shaped by social factors, contributing to our understanding of the evolution of human nonverbal communication, language, and tool use.

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.

Don't look back on failure: spontaneous uncertainty monitoring in chimpanzees.

During computer-controlled cognitive tasks, chimpanzees often look up at the food dispenser, which activates at the same time as feedback for the correct choice but not for feedback for the incorrect choice. Do these "looking back" behaviors also indicate signs of spontaneous monitoring of their confidence in their choices? To address this question, we delayed the feedback for 1 s after their choice responses and observed their look-back behaviors during the delay period. Two chimpanzees looked up at the food dispenser significantly less frequently when their choice was incorrect (but the feedback was not given) than when it was correct. These look-back behaviors have not been explicitly trained under experimental contexts. Therefore, these results indicate that chimpanzees spontaneously change the frequency of their look-back behaviors in response to the correctness or incorrectness of their own choices, even without external feedback, suggesting that their look-back behaviors may reflect the level of "confidence" or "uncertainty" of their responses immediately before.

I'll (not) take that: The reverse-reward contingency task as a test of self-control and inhibition.

While searching for more evidence of quantitative skills in chimpanzees to add to what she already had found, Boysen discovered something else. When training chimpanzees to point at what they would not get, and not pointing at what they would get, none could do this for piles of food items. Even when those items in the pointed-at set were given away to another chimpanzee, and even with experience in the task, failure persisted. This test, the reverse-reward contingency test, has now been used with many species, as a means of assessing inhibitory control and perhaps self-control in animals. Typically, the task is difficult, and only specific manipulations have worked to allow primates to overcome the reversed contingencies. This includes using symbolic stimuli, adding another layer to the story, and more value to the task itself as a measure perhaps of forms of cognitive control in other species. I will discuss some of these empirical results, including from other chimpanzees who were given variations of the task, and how these studies have influenced numerous areas within comparative cognitive science.

Attention to inattention: Effect of the other's attentional state on visual search performance in chimpanzees.

In the present study, we examined the effects of the other's triadic attention to objects on visual search performances in chimpanzees. We found the search-asymmetry-like effect of the other's attentional state; the chimpanzees searched a target object not attended by the other individual more efficiently than that attended (Experiment 1). Additional experiments explored the possibility that the other individual "holding an object but not looking at it" led to expectancy violation (Experiment 2) or the role of nonsocial cues such as the proximity relation between the head and the object (Experiment 3). Still, these accounts alone did not explain this effect. It was also shown that the other's attentional state affected the chimpanzees' performances more readily as the interference effect than the facilitation effect (Experiment 4). Furthermore, the same effect was observed in the visual search for the gaze (head direction) of others (Experiment 5). We obtained the same results using photographs of chimpanzees (Experiment 6). Contrary to the chimpanzees, humans detected the object to which attention was directed more efficiently than vice versa (Experiment 7). The present results may reflect species differences between chimpanzees and humans in processing triadic social attention.

Decay rates of arboreal and terrestrial nests of Eastern chimpanzees (Pan troglodytes schweinfurthii) in the Bugoma Central Forest Reserve, Uganda: Implications for population size estimates.

Chimpanzees were once thought to sleep primarily in the trees, but recent studies indicate that some populations also construct terrestrial night nests. This behavior has relevance not only to understanding the behavioral diversity of Pan troglodytes, but also to the conservation of the species, given that nest encounter rates are often used to estimate great ape population densities. A proper estimate of decay rates for ground nests is necessary for converting the encounter rate of nests to the density of weaned chimpanzees. Here we present the results of the first systematic comparative study between the decay rates of arboreal and terrestrial chimpanzee nests, from the Bugoma Central Forest Reserve in western Uganda. We followed the decay of 56 ground and 51 tree nests in eight nest groups between April 2020 and October 2021. For 15 of the ground and 19 of the tree nests, we collected detailed information on the condition of the nests every two weeks; we checked the remaining 73 nests only twice. On average, ground nests lasted 238 days versus 276 days for tree nests (p = 0.05). Of the 107 total nests surveyed, 51% of tree and 64% of ground nests had disappeared after six months. Based on our results, we propose a modification of the formula used to convert nest density into chimpanzee density. Our results highlight the importance of taking into account potential differences in decay rates between ground versus tree nests, which will likely influence our understanding of the distribution of ground nesting behavior in chimpanzee across tropical Africa, as well as our estimations of the densities of ground nesting populations.

Rhythmic swaying induced by sound in chimpanzees (Pan troglodytes).

Music and dance are universal across human culture and have an ancient history. One characteristic of music is its strong influence on movement. For example, an auditory beat induces rhythmic movement with positive emotions in humans from early developmental stages. In this study, we investigated if sound induced spontaneous rhythmic movement in chimpanzees. Three experiments showed that: 1) an auditory beat induced rhythmic swaying and other rhythmic movements, with larger responses from male chimpanzees than female chimpanzees; 2) random beat as well as regular beat induced rhythmic swaying and beat tempo affected movement periodicity in a chimpanzee in a bipedal posture; and 3) a chimpanzee showed close proximity to the sound source while hearing auditory stimuli. The finding that male chimpanzees showed a larger response to sound than female chimpanzees was consistent with previous literature about "rain dances" in the wild, where male chimpanzees engage in rhythmic displays when hearing the sound of rain starting. The fact that rhythmic swaying was induced regardless of beat regularity may be a critical difference from humans, and a further study should reveal the physiological properties of sound that induce rhythmic movements in chimpanzees. These results suggest some biological foundation for dancing existed in the common ancestor of humans and chimpanzees ∼6 million years ago. As such, this study supports the evolutionary origins of musicality.

Oxytocin promotes species-relevant outgroup attention in bonobos and chimpanzees.

Previous research has found that oxytocin (OT) is associated with intergroup behaviour in humans as well as wild chimpanzees, and that exogenous OT affects Pan social attention. The two Pan species, bonobos and chimpanzees, differ drastically from one another in their intensity of intergroup competition, with lethal intergroup aggression often led by males in chimpanzees and more tolerant associations often centered around females in bonobos. However, it remains unclear how exogenous OT changes the two species' responses to ingroup and outgroup individuals. In this study, after intranasal administration of nebulized OT or placebo control, chimpanzees and bonobos viewed image pairs of ingroup and outgroup conspecifics while their eye movements were tracked with an eye-tracker. Although the overall effect of OT was small, we found that OT shifted bonobos' and chimpanzees' attention to outgroup images of the sex primarily involved in intergroup encounters in each species. Specifically, OT selectively shifted attention towards outgroup photos of female conspecifics in bonobos, and those of outgroup male conspecifics in chimpanzees. This suggests that OT generally promotes outgroup attention in both bonobos and chimpanzees but this effect is restricted to the sex most relevant in intergroup relations. These results suggest that, although OT may have a generally conserved role in hominid intergroup behaviour, it may act in species-relevant ways under the influence of their socio-ecological backgrounds.

Chimpanzees (Pan troglodytes) detect strange body parts: an eye-tracking study.

This study investigated chimpanzee body representation by testing whether chimpanzees detect strangeness in body parts. We tested six chimpanzees with edited chimpanzee body pictures in eye-tracking tasks. The target body parts were arms or legs. For either target, there were four conditions: "normal" condition as control, where all bodies were normal; "misplaced" condition, where one arm or one leg was misplaced to an incorrect body location in each picture; "replaced by a chimpanzee part" condition, where one arm or one leg was replaced by a chimpanzee leg or arm, respectively, in its original place in each picture; and "replaced by a human part" condition, where one arm or one leg was replaced by a human arm or leg in each picture. Compared to the looking times toward the normal parts, chimpanzees had significantly longer looking times toward the human arms or legs. The looking times toward the misplaced parts were also longer than the normal parts, but the difference just failed to meet significance. These results indicate more interests toward strange body parts, compared to typical parts, suggesting that chimpanzees might have a body representation that is sufficiently sensitive to detect these aspects of strangeness.

Chimpanzees consider freedom of choice in their evaluation of social action.

Judgements of wrongdoing in humans often hinge upon an assessment of whether a perpetrator acted out of free choice: whether they had more than one option. The classic inhibitors of free choice are constraint (e.g. having your hands tied together) and ignorance (e.g. being unaware that an alternative exists). Here, across two studies, we investigate whether chimpanzees consider these factors in their evaluation of social action. Chimpanzees interacted with a human experimenter who handed them a non-preferred item of food, either because they were physically constrained from accessing the preferred item (Experiment 1) or because they were ignorant of the availability of the preferred item (Experiment 2). We found that chimpanzees were more likely to accept the non-preferred food and showed fewer negative emotional responses when the experimenter was physically constrained compared with when they had free choice. We did not, however, find an effect of ignorance on chimpanzee's evaluation. Freedom of choice factors into chimpanzees' evaluation of how they are treated, but it is unclear whether mental state reasoning is involved in this assessment.

Sulcal Morphology in Cingulate Cortex is Associated with Voluntary Oro-Facial Motor Control and Gestural Communication in Chimpanzees (Pan troglodytes).

Individual differences in sulcal variation within the anterior and mid-cingulate cortex of the human brain, particularly the presence or absence of a paracingulate sulcus (PCGS), are associated with various motor and cognitive processes. Recently, it has been reported that chimpanzees possess a PCGS, previously thought to be a unique feature of the human brain. Here, we examined whether individual variation in the presence or absence of a PCGS as well as the variability in the intralimbic sulcus (ILS) are associated with oro-facial motor control, handedness for manual gestures, and sex in a sample of MRI scans obtained in 225 chimpanzees. Additionally, we quantified the depth of the cingulate sulcus (CGS) along the anterior-posterior axis and tested for association with oro-facial motor control, handedness, and sex. Chimpanzees with better oro-facial motor control were more likely to have a PCGS, particularly in the left hemisphere compared to those with poorer control. Male chimpanzees with better oro-facial motor control showed increased leftward asymmetries in the depth of the anterior CGS, whereas female chimpanzees showed the opposite pattern. Significantly, more chimpanzees had an ILS in the left compared to the right hemisphere, but variability in this fold was not associated with sex, handedness, or oro-facial motor control. Finally, significant population-level leftward asymmetries were found in the anterior portion of the CGS, whereas significant rightward biases were evident in the posterior regions. The collective results suggest that the emergence of a PCGS and enhanced gyrification within the anterior and mid-cingulate gyrus may have directly or indirectly evolved in response to selection for increasing oro-facial motor control in primates.

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.

Male chimpanzee sexual coercion and mating success at Ngogo.

Chimpanzees (Pan troglodytes) have a complex mating system in which both sexes use multiple tactics. Most copulations occur in group contexts, and high-ranking males can gain high mating and reproductive success, but females typically mate with multiple males and the potential for sperm competition is high. Also, male-female dyads sometimes form temporary exclusive mating associations (consortships). Male aggression to receptive females is common. Several studies have supported the hypothesis that this is sexual coercion, but debate exists regarding the importance of coercion relative to that of female choice. The number of adult males in a community can influence the balance between these processes. In the large Ngogo community, male dominance ranks and rates of aggression to fully-swollen females were positively related to mating success as estimated by copulation rates and by proportions of copulations achieved. Aggression rates were higher than at other sites, overall and per male, especially during periovulatory periods, and increased with the number of males associating with a female. Aggression impaired female foraging efficiency. Males initiated most copulations and females rarely refused mating attempts. Male-to-female grooming was positively associated with male mating success and with the proportion of copulations that females initiated, but the amount of grooming was typically small and whether grooming-for-mating trading occurs is uncertain. These results are consistent with the hypothesis that sexual coercion is an important component of male chimpanzee mating strategies in many sociodemographic circumstances, but also show that male tactics vary both in response to and independently of those circumstances.