Searching for the earliest archaeological record: insights from chimpanzee material landscapes.

The origin of tool use is a central question in human evolutionary studies. Plio-Pleistocene core and flake technologies represent the earliest evidence of tool use in the human lineage. Some suggest this form of tool use is probably pre-dated by a phase of percussive tool use. However, there is currently no evidence for such a record. The archaeological signature of solely percussive behaviours is not as well understood as that associated with cores and flakes. The durable nature of primate percussive stone tools and their by-products provide an opportunity to investigate what such a record looks like. Here, we present a landscape-scale study of the chimpanzee (Pan troglodytes verus) material culture from the Djouroutou Chimpanzee Project, Taï Forest, Cote d'Ivoire. This study explores the interplay between behavioural and environmental factors in shaping the stone record of nut cracking. Through a survey of nut-cracking sites, the available nut species, and raw materials, we show how resource availability influences the resulting material signature of nut cracking. These results also reveal the diversity of material signatures associated with a purely percussive material record. We gain insight into the range of signatures that may be associated with a pre-core and flake archaeological record, providing new expectations for an earlier record of tool use.

Body part categorical matching in chimpanzees (Pan troglodytes).

Humans categorize body parts, reflecting our knowledge about bodies, and this could be useful in higher-level activities involving bodies. We tested whether humans' closest living relatives-chimpanzees-have the same ability using touchscreen tasks, focusing on the major parts: heads, torsos, arms, and legs. Six chimpanzees were trained to perform a body part matching-to-sample task using sets of pictures of chimpanzee bodies, where in each trial, the sample and choice pictures were the same. Five passed the training and received the test sessions, where three trial types were mixed: trained same-individual picture pairs; novel same-individual picture pairs; and novel different-individual picture pairs. All participants performed better than the chance level in all conditions and for all body parts. Further analyses showed differences in performance when the samples were different body parts. For example, the results revealed better performances for heads and torsos than arms and legs in "novel different-individual pairs". The study showed that chimpanzees can visually match and categorize body parts in this experiment setting, even across different chimpanzees' bodies, suggesting potential biological understanding. Different performances for body parts suggested a deviated categorization from humans. We hope this study will inspire future research on the evolution of body perception.

Treponematosis in critically endangered Western chimpanzees (Pan troglodytes verus) in Senegal.

Treponematoses encompass a group of chronic and debilitating bacterial diseases transmitted sexually or by direct contact and attributed to Treponema pallidum. Despite being documented since as far back as 1963, the epidemiology of treponematoses in wild primates has remained an uninvestigated territory due to the inherent challenges associated with conducting examinations and obtaining invasive biological samples from wild animals. The primary aim of this study was to investigate the presence of treponemal infections in the critically endangered Western chimpanzees in Senegal, utilizing an innovative non-invasive stool serology method. We provide compelling evidence of the existence of anti-Treponema-specific antibodies in 13 out of 29 individual chimpanzees. Our study also underscores the significant potential of stool serology as a valuable non-invasive tool for monitoring and surveilling crucial emerging diseases in wild animals. We recognize two major implications: (1) the imperative need to assess the risks of treponematosis in Western chimpanzee populations and (2) the necessity to monitor and manage this disease following a holistic One Health approach.

Strontium isotopes track female dispersal in Taï chimpanzees.

OBJECTIVES: Chimpanzees (Pan troglodytes) are patrilocal, with males remaining in their natal community and females dispersing when they reach sexual maturity. However, the details of female chimpanzee dispersal, such as their possible origin, are difficult to assess, even in habituated communities. This study investigates the utility of 87Sr/86Sr analysis for (1) assessing Sr baseline differences between chimpanzee territories and (2) identifying the status (immigrant or natal) of females of unknown origin within the territories of five neighboring communities in Taï National Park (Côte d'Ivoire). MATERIALS AND METHODS: To create a local Sr isoscape for the Taï Chimpanzee Project (TCP) study area, we sampled environmental samples from TCP-established territories (n = 35). To assess dispersal patterns, 34 tooth enamel samples (one per individual) were selected from the Taï chimpanzee skeletal collection. 87Sr/86Sr analysis was performed on all 69 samples at the W.M. Keck Lab. The theoretical density and overlap of chimpanzee communities as well as generalized linear mixed models (GLMMs) were used to test each question. RESULTS: 87Sr/86Sr ratios for natal male chimpanzees ranged from 0.71662 to 0.72187, which is well within the corresponding environmental baseline range of 0.70774-0.73460. The local Sr isoscapes fit was estimated with the root-mean-square error value, which was 0.0048 (22% of the whole 87Sr/86Sr data range). GLMMs identified significant differences in 87Sr/86Sr ratios between natal and unknown North community origin groups, suggesting that after 1980, females of unknown origin could be immigrants to North community (n = 7, z-ratio = -4.08, p = 0.0001, power = 0.94). DISCUSSION: This study indicates that 87Sr/86This study indicates that 87Sr/86Sr analysis can successfully identify immigrant females in skeletal collections obtained from wild chimpanzee communities, enabling the tracking of female dispersal patterns historically. There are, however, significant limitations within the scope of this study, such as (1) the absence of reliable maps for the TCP study area, (2) limited capacity for environmental sampling, (3) small sample sizes, and (4) tooth formation in wild chimpanzees.

Sodium content in plant and insect food resources consumed by chimpanzees (Pan troglodytes schweinfurthii) in Gombe National Park, Tanzania.

OBJECTIVES: Many nonhuman primate diets are dominated by plant foods, yet plant tissues are often poor sources of sodium-a necessary mineral for metabolism and health. Among primates, chimpanzees (Pan troglodytes), which are ripe fruit specialists, consume diverse animal, and plant resources. Insects have been proposed as a source of dietary sodium for chimpanzees, yet published data on sodium values for specific foods are limited. We assayed plants and insects commonly eaten by chimpanzees to assess their relative value as sodium sources. MATERIALS AND METHODS: We used atomic absorption spectroscopy to determine sodium content of key plant foods and insects consumed by chimpanzees of Gombe National Park, Tanzania. Dietary contributions of plant and insect foods were calculated using feeding observational data. RESULTS: On a dry matter basis, mean sodium value of plant foods (n = 83 samples; mean = 86 ppm, SD = 92 ppm) was significantly lower than insects (n = 12; mean = 1549 ppm, SD = 807 ppm) (Wilcoxon rank sum test: W = 975, p < 0.001). All plant values were below the suggested sodium requirement (2000 ppm) for captive primates. While values of assayed insects were variable, sodium content of two commonly consumed insect prey for Gombe chimpanzees (Macrotermes soldiers and Dorylus ants) were four to five times greater than the highest plant values and likely meet requirements. DISCUSSION: We conclude that plant foods available to Gombe chimpanzees are generally poor sources of sodium while insects are important, perhaps critical, sources of sodium for this population.

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.

Inconsistent effects of components as evidence for non-compositionality in chimpanzee face-gesture combinations? A response to Oña et al (2019).

Using field observations from a sanctuary, Oña and colleagues (DOI: 10.7717/peerj.7623) investigated the semantics of face-gesture combinations in chimpanzees (Pan troglodytes). The response of the animals to these signals was encoded as a binary measure: positive interactions such as approaching or grooming were considered affiliative; ignoring or attacking was considered non-affiliative. The relevant signals are illustrated in Fig. 1 (https://doi.org/10.7717/peerj.7623/fig-1), together with the outcome in terms of average affiliativeness. The authors observe that there seems to be no systematicity in the way the faces modify the responses to the gestures, sometimes reducing affiliativeness, sometimes increasing it. A strong interpretation of this result would be that the meaning of a gesture-face combination cannot be derived from the meaning of the gesture and the meaning of the face, that is, the interpretation of chimpanzees' face-gesture combinations are non compositional in nature. We will revisit this conclusion: we will exhibit simple compositional systems which, after all, may be plausible. At the methodological level, we argue that it is critical to lay out the theoretical options explicitly for a complete comparison of their pros and cons.

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.

Chimpanzees engage in competitive altruism in a triadic ultimatum game.

Partner choice promotes competition among individuals to be selected as a cooperative partner, a phenomenon referred to as competitive altruism. We explored whether chimpanzees engage in competitive altruism in a triadic Ultimatum Game where two proposers can send offers simultaneously or consecutively to a responder who can only accept one of the two competing offers. In a dyadic control condition only one proposer at a time could send an offer to the responder. Chimpanzees increased their offers across trials in the competitive triadic, but not in the dyadic control condition. Chimpanzees also increased their offers after being rejected in previous triadic trials. Furthermore, we found that chimpanzees, under specific conditions, outcompete first proposers in triadic consecutive trials before the responder could choose which offer to accept by offering more than what is expected if they acted randomly or simply offered the smallest possible amount. These results suggest that competitive altruism in chimpanzees did not emerge just as a by-product of them trying to increase over previous losses. Chimpanzees might consider how others' interactions affect their outcomes and engage in strategies to maximize their chances of being selected as cooperative partners.

May the force be with you: exploring force discrimination in chimpanzees using the force-feedback device.

While force-feedback devices have been developed in areas such as virtual reality, there have been very few comparative cognitive studies in nonhuman animals using these devices. In addition, although cross-modal perception between vision and touch has been actively studied in nonhuman primates for several decades, there have been no studies of their active haptic perception. In this study, we attempted to train force discrimination in chimpanzees using a force-feedback device modified from a trackball. Chimpanzees were given different levels of force feedback (8.0 vs. 0.5 N) when moving the on-screen cursor to the target area by manipulating the trackball and were required to select one of two choice stimuli based on the force cue. The experiment was conducted using a trial-block procedure in which the same force stimulus was presented for a fixed number of trials, and the force stimulus was changed between blocks. The block size was progressively reduced from ten trials. Four chimpanzees were trained, but none reached the learning criterion (80% or more correct responses under the condition that the force stimuli were presented randomly). However, a detailed analysis of the chimpanzees' performance before and after the trial-block switching revealed that their choice behavior could not be explained by a simple win-stay/lose-shift strategy, suggesting that the switching of the force stimuli affected the chimpanzees' choice behavior. It was also found that the chimpanzees performed better when switching from small to large force stimuli than when switching from large to small force stimuli. Although none of the chimpanzees in this study acquired force discrimination, future studies using such force-feedback devices will provide new insights for understanding haptic cognition in nonhuman primates from a comparative cognitive perspective.

Female chimpanzees avoid inbreeding even in the presence of substantial bisexual philopatry.

Inbreeding (reproduction between relatives) often decreases the fitness of offspring and is thus expected to lead to the evolution of inbreeding avoidance strategies. Chimpanzees (Pan troglodytes) are expected to avoid inbreeding as they are long-lived, invest heavily in offspring and may encounter adult, opposite sex kin frequently, especially in populations where both males and females commonly remain in the group in which they were born (bisexual philopatry). However, it is unclear whether substantial bisexual philopatry has been a feature of chimpanzees' evolutionary history or whether it is a result of recent anthropogenic interference, as the only groups for which it has been documented are significantly impacted by human encroachment and experience notable rates of potentially unsustainable inbreeding. Here we use 14 years of observational data and a large genomic dataset of 256 481 loci sequenced from 459 individuals to document dispersal and inbreeding dynamics in an eastern chimpanzee (P. t. schweinfurthii) community with low levels of anthropogenic disturbance. We document the first case of substantial bisexual philopatry in a relatively undisturbed chimpanzee community and show that, despite an increased inbreeding risk incurred by females who do not disperse before reaching reproductive age, natal females were still able to avoid producing inbred offspring.

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.

Characterizing the personality and gray matter volume of chimpanzees that exhibit autism-related socio-communicative phenotypes.

Autism spectrum disorder (ASD) is a developmental disorder characterized by stereotypies or repetitive behaviors and impairments in social behavior and socio-communicative skills. One hallmark phenotype of ASD is poor joint attention skills compared to neurotypical controls. In addition, individuals with ASD have lower scores on several of the Big 5 personality dimensions, including Extraversion. Here, we examine these traits in a nonhuman primate model (chimpanzees; Pan troglodytes) to further understand the relationship between personality and joint attention skills, as well as the genetic and neural systems that contribute to these phenotypes. We used archival data including receptive joint attention (RJA) performance, personality based on caretaker ratings, and magnetic resonance images from 189 chimpanzees. We found that, like humans, chimpanzees who performed worse on the RJA task had lower Extraversion scores. We also found that joint attention skills and several personality dimensions, including Extraversion, were significantly heritable. There was also a borderline significant genetic correlation between RJA and Extraversion. A conjunction analysis examining gray matter volume showed that there were five main brain regions associated with both higher levels of Extraversion and social cognition. These regions included the right posterior middle and superior temporal gyrus, bilateral inferior frontal gyrus, left inferior frontal sulcus, and left superior frontal sulcus, all regions within the social brain network. Altogether, these findings provide further evidence that chimpanzees serve as an excellent model for understanding the mechanisms underlying social impairment related to ASD. Future research should further examine the relationship between social cognition, personality, genetics, and neuroanatomy and function in nonhuman primate models.

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.

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.

Assessing Eysenck's PEN model to describe personality in chimpanzees.

Questionnaires based on human models can be used to reliably assess personality also in non-human primates. In this study, we used an adapted version of Eysenck's Psychoticism-Extraversion-Neuroticism (PEN) model that focuses on three higher-order personality traits. Extending previous work on a small group of chimpanzees (Pan troglodytes), we tested 37 chimpanzees housed at Fundació Mona (Girona, Spain) and the Leipzig Zoo (Germany). We assessed personality with a 12-item questionnaire, which raters scored using a 7-point Likert scale. To identify the personality traits, we conducted data reduction with Principal Components Analysis and Robust Unweighted Least Squares. The ICCs for the single (3, 1) and average (3, k) ratings indicated substantial agreement between raters. Parallel analyses identified two factors to retain, whereas the scree plot inspection and eigenvalues larger than one rule identified three factors. Factor 1 and 2 in our study were identical to the ones previously described for this species (labelled Extraversion and Neuropsychoticism, respectively) and we also obtained a third factor that could be related to Dominance (Fearless Dominance). Thus, our results confirm the potential of the PEN model to describe chimpanzee personality structure.

The impact of sanctuary visits on children's knowledge and attitudes toward primate welfare and conservation.

Primate sanctuaries provide a solution for the increasing number of primates being taken from their home countries to support the demands of the illegal pet trade. To help end the primate trade and raise awareness about the risks this trade poses to delicate ecosystems, sanctuaries are increasingly developing conservation education programs. Education and raising awareness must be one of the primary roles of primate sanctuaries. However, there are few evaluations of the impacts of conservation education programs for school children published in scientific literature. To address this gap, we conducted an evaluation of educator-led visits of school children at Fundació Mona, a primate sanctuary located in Catalunya, Spain. Questionnaires for an experimental and control group were conducted with 3,205 school children, ages 8 to 18 from 83 different schools, to evaluate changes in their attitudes and knowledge of primate welfare and conservation. We found that Fundació Mona's program of environmental activities had a positive impact on children, both female and male students, in terms of attitudes and knowledge of primate welfare and conservation. Although female students gave better responses regarding welfare and conservation, all children showed gains in pro-conservation responses. This study demonstrates that environmental education activities focused on children can help shape a change in knowledge and attitudes toward primate welfare and conservation. Educator-led visits of school children to primate sanctuaries such as Fundació Mona can also serve to amplify biodiversity conservation messages among children and their families. We encourage primate sanctuaries to promote empirical studies of attitudes and knowledge of primate welfare and conservation and to conduct systematical evaluations to strengthen their educational activities.

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.