Captive Chimpanzees Research Articles

Long-Term Assessment of Captive Chimpanzees: Influence of Social Group Composition, Seasonality and Biographic Background.

Wild chimpanzees (Pan troglodytes) live in flexible fission-fusion societies with frequent changes in both group size and composition. These changes depend mostly on resource availability and individual social preferences yet in captivity are determined by housing organizations. During a period of seven years, we observed a group of sanctuary chimpanzees, focusing on how they adapted to changes in the group composition over time. Using linear mixed models (LMMs), factors such as group size, sex ratio, seasonality, and the individuals' sex and origin (wild caught vs. captive born) were considered in order to evaluate the impact on the chimpanzees' activity levels, the occurrence of undesired behaviors (abnormal and self-directed behaviors) and the social grooming networks. Our results indicate that the activity levels and the occurrence of undesired behaviors were impacted by changes in group composition and the individuals' biographic background. The colder season was marked by higher levels of activity and more time spent grooming. Moreover, it was the individuals' origin but not changes in group composition that affected social grooming, with wild-caught chimpanzees grooming far less frequently. Long-term observations are essential to evaluate, predict and detect potential benefits and/or issues of housing conditions while considering the social and physical environment.

Does tolerance allow bonobos to outperform chimpanzees on a cooperative task? A conceptual replication of Hare et al., 2007.

Across various taxa, social tolerance is thought to facilitate cooperation, and many species are treated as having species-specific patterns of social tolerance. Yet studies that assess wild and captive bonobos and chimpanzees result in contrasting findings. By replicating a cornerstone experimental study on tolerance and cooperation in bonobos and chimpanzees (Hare et al. 2007 Cur. Biol. 17, 619-623 (doi:10.1016/j.cub.2007.02.040)), we aim to further our understanding of current discrepant findings. We tested bonobos and chimpanzees housed at the same facility in a co-feeding and cooperation task. Food was placed on dishes located on both ends or in the middle of a platform. In the co-feeding task, the tray was simply made available to the ape duos, whereas in the cooperation task the apes had to simultaneously pull at both ends of a rope attached to the platform to retrieve the food. By contrast to the published findings, bonobos and chimpanzees co-fed to a similar degree, indicating a similar level of tolerance. However, bonobos cooperated more than chimpanzees when the food was monopolizable, which replicates the original study. Our findings call into question the interpretation that at the species level bonobos cooperate to a higher degree because they are inherently more tolerant.

A Preliminary Assessment of Compassion Fatigue in Chimpanzee Caregivers.

Compassion fatigue is defined as "traumatization of helpers through their efforts at helping others". It has negative effects on clinicians including reduced satisfaction with work, fatigue, irritability, dread of going to work, and lack of joy in life. It is correlated with patients' decreased satisfaction with care. Compassion fatigue occurs in a variety of helping professions including educators, social workers, mental health clinicians, and it also appears in nonhuman animal care workers. This study surveyed caregivers of chimpanzees using the ProQOL-V to assess the prevalence of compassion fatigue among this group. Compassion satisfaction is higher than many other types of animal care workers. Conversely, this group shows moderate levels of burnout and secondary traumatic stress; higher levels than other types of animal care workers and many medical professions. While compassion fatigue has an effect on the caregiver's experience, it has potential to affect animal welfare. Caregivers are an integral part of the chimpanzee social network. Compassion fatigue affects the caregiver's attitude, this could in turn affect the relationship and degrade the experience of care for captive chimpanzees. Compassion fatigue can be mitigated with professional development, mindfulness training, interrelationships among staff, and specialized training. This preliminary assessment indicates the work ahead is educating caregivers about compassion fatigue and implementing procedures in sanctuaries to mitigate burnout and secondary traumatic stress.

Wild Chimpanzee Welfare: A Focus on Nutrition, Foraging and Health to Inform Great Ape Welfare in the Wild and in Captivity

Adequate nutrition is essential for individual well-being, survival and reproductive fitness. Yet, in wild animals, including great apes, scoring nutrition or health comes with many challenges. Here, we have two aims: first, broadly review the scientific literature regarding nutritional data on wild chimpanzee foods to get a better understanding what nutrients foods comprise of, and second, highlight important findings on wild chimpanzee nutrition and welfare pertaining to diet. We discuss variation in macro and micronutrients in food items consumed and their role in chimpanzee health across chimpanzee subspecies from multiple study sites. We found a lack of information pertaining to nutritional consumption rates of daily diets. Second, we call for a fresh, in-depth discussion on wild chimpanzee welfare issues is of foremost importance to inform conservation projects and particularly settings where humans and chimpanzees may interact, because such conversation can reveal how specific or general welfare measures can (a) inform our knowledge of an individual's, group's, and population's welfare, (b) provide additional measures from the study of wild chimpanzee ecology that can guide the welfare of captive chimpanzees, and (c) can enable comparative study of welfare across wild populations. A summary of the current literature on approaches to measuring wild chimpanzee health and welfare status, to our knowledge, has yet to be done.

Neuroanatomical correlates of individual differences in the object choice task in chimpanzees (Pan troglodytes).

Declarative and imperative joint attention or joint engagement are important milestones in human infant development. These have been shown to be a significant predictor of later language development and are impaired in some individuals with, or at risk for, a diagnosis of autism spectrum disorder. Comparatively, while chimpanzees and other great apes have been reported to engage in imperative joint attention, evidence of declarative joint attention remains unclear based on existing studies. Some have suggested that differences in methods of assessing joint attention may have an influence on performance in nonhuman primates. Here, we report data on a measure of receptive joint attention (object choice task) in a sample of captive chimpanzees. Chimpanzees, as a group, performed significantly better than chance. By contrast, when considering individual performance, there was no significant difference in the number of those who passed and those who failed. Using quantitative genetic analyses, we found that performance on the object choice task was not significantly heritable nor were there any significant effects of sex, rearing history, or colony. Lastly, we found significant differences in gray matter covariation, between those who passed or failed the task. Those who passed contributed more to gray matter covariation in several brain regions within the social brain network, consistent with hypotheses regarding the importance of these regions in human and nonhuman primate social cognition.

Bone-related behaviours of captive chimpanzees (Pan troglodytes) during two excavating experiments

After stone tools, bone tools are the most abundant artefact type in the Early Pleistocene archaeological record. That said, they are still relatively scarce, which limits our understanding of the behaviours that led to their production and use. Observations of extant primates constitute a unique source of behavioural data with which to construct hypotheses about the technological forms and repertoires exhibited by our hominin ancestors. We conducted two different experiments to investigate the behavioural responses of two groups of captive chimpanzees (Pan troglodytes; n = 33 and n = 9) to disarticulated, defleshed, ungulate bones while participating in a foraging task aimed at eliciting excavating behaviour. Each chimpanzee group was provided with bone specimens with different characteristics, and the two groups differed in their respective experience levels with excavating plant tools. We found that several individuals from the inexperienced group used the provided bones as tools during the task. In contrast, none of the individuals from the experienced group used bones as excavating tools, but instead continued using plant tools. These chimpanzees also performed non-excavating bone behaviours such as percussion and tool-assisted extraction of organic material from the medullary cavity. Our findings serve as a proof-of-concept that chimpanzees can be used to investigate spontaneous bone tool behaviours such as bone-assisted excavation. Furthermore, our results raise interesting questions regarding the role that bone characteristics, as well as previous tool-assisted excavating experience with other raw materials, might have in the expression of bone tool-assisted excavation.

The Role of Serotonergic Gene Methylation in Regulating Anxiety-Related Personality Traits in Chimpanzees.

While low serotonergic activity is often associated with psychological disorders such as depression, anxiety, mood, and personality disorders, variations in serotonin also contribute to normal personality differences. In this study, we investigated the role of blood DNA methylation levels at individual CpG sites of two key serotonergic genes (serotonin receptor gene 1A, HTR1A; serotonin transporter gene, SLC6A4) in predicting the personalities of captive chimpanzees. We found associations between methylation at 9/48 CpG sites with four personality dimensions: Dominance, Reactivity/Dependability, Agreeableness, and Openness. Directionality of effects were CpG location-dependent and confirmed a role of serotonergic methylation in reducing anxiety (Dominance) and aggression-related personality (Reactivity/Undependability) while simultaneously promoting prosocial (Agreeableness) and exploratory personalities (Openness). Although early-life adversity has been shown to impact serotonergic methylation patterns in other species, here, atypical early social rearing experiences only had a modest impact on CpG methylation levels in this chimpanzee sample. The precise environmental factors impacting serotonergic methylation in chimpanzees remain to be identified. Nevertheless, our study suggests a role in shaping natural variation in animal personalities. The results of this study offer a basis for future hypothesis-driven testing in additional populations and species to better understand the impact of ecology and evolution on complex behavioral traits.

Age, adrenal steroids, and cognitive functioning in captive chimpanzees (Pan troglodytes).

Dehydroepiandrosterone-sulfate is the most abundant circulating androgen in humans and other catarrhines. It is involved in several biological functions, such as testosterone production, glucocorticoid antagonist actions, neurogenesis and neuroplasticty. Although the role of dehydroepiandrosterone-sulfate (DHEAS) in cognition remains elusive, the DHEAS/cortisol ratio has been positively associated with a slower cognitive age-decline and improved mood in humans. Whether this relationship is found in nonhuman primates remains unknown. We measured DHEAS and cortisol levels in serum of 107 adult chimpanzees to investigate the relationship between DHEAS levels and age. A subset of 21 chimpanzees was used to test the potential associations between DHEAS, cortisol, and DHEAS/cortisol ratio in cognitive function, taking into account age, sex, and their interactions. We tested for cognitive function using the primate cognitive test battery (PCTB) and principal component analyses to categorize cognition into three components: spatial relationship tasks, tool use and social communication tasks, and auditory-visual sensory perception tasks. DHEAS levels, but not the DHEAS/cortisol ratio, declined with age in chimpanzees. Our analyses for spatial relationships tasks revealed a significant, positive correlation with the DHEAS/cortisol ratio. Tool use and social communication had a negative relationship with age. Our data show that the DHEAS/cortisol ratio, but not DHEAS individually, is a promising predictor of spatial cognition in chimpanzees.

The Association Between the Bared-Teeth Display and Social Dominance in Captive Chimpanzees (Pan troglodytes)

Humans use smiles — widely observed emotional expressions — in a variety of social situations, of which the meaning varies depending on social relationship and the context in which it is displayed. The homologue of the human smile in non-human primates — both due to morphological and functional similarities — is the bared-teeth display (BT). According to the power asymmetry hypothesis (PAH), species with strict linear dominance hierarchies are predicted to produce distinct communicative signals to avoid escalations of social conflicts. Hence, while the BT in a despotic species is predicted to be expressed from low- to high-ranking individuals, signaling submission, the BT in a tolerant species is predicted to be expressed in multiple contexts, regardless of rank. We tested this hypothesis in a group of 8 captive chimpanzees (Pan troglodytes), a species commonly characterized as rather despotic. An investigation of 11,774 dyadic social interactions revealed this chimpanzee group to have a linear dominance hierarchy, with moderate steepness. A Bayesian GLMM — used to test the effects of social contexts and rank relationships of dyads on the use of the BT display — indicated multi-contextual use of the BT which is contingent on the rank relationship. We also found that slight morphological and/or acoustic variants (i.e., silent bared-teeth and vocalized bared-teeth) of the BT display may have different communicative meanings. Our findings are in line with the prediction derived from the PAH for a moderately despotic species, and the view that the human smile originated from the primate BT display.

Long-Term Spatial Restriction Generates Deferred Limited Space Use in a Zoo-Housed Chimpanzee Group.

Simple SummaryA worldwide trend amongst zoos is to replacesmall, barren enclosures with large, naturalistic ones intended to provide animals with environments which cater to their behavioural and psychological needs. Evidence suggests that naturalistic enclosures are effective but most studies focus on welfare-related behaviour or human perceptions of the enclosures. To date, little attention has been given to how animals use space in naturalistic enclosures. Our study investigated how a group of chimpanzees at the Johannesburg Zoo used space in a naturalistic enclosure by recording behaviour and space use every 5 min for an hour at a time. We found that the chimpanzees showed a preference for locations within the enclosure which coincided with their previous housing and that the chimpanzees form subgroups which conform to the space of their previous housing (i.e., small, barren enclosure). We suggest that the chimpanzees’ perception of space has been altered by their experience of the previous, smaller barren housing and that this limits their space use in the naturalistic enclosure through what appears to be a self-imposed ‘invisible cage’ around subgroups. Exactly how the ‘invisible cage’ works is unclear but our findings have implications for animal welfare, husbandry and broader conservation of endangered species.Background: Appropriate space is considered paramount for good captive animal welfare. There has been a concerted effort by captive institutions, particularly zoos, to provide captive animals with relatively large, naturalistic enclosures which havehad demonstrated welfare benefits for animals. However, post-occupancy assessments of these enclosures tend to focus on short-term welfare-centredbehavioural effects or human perceptions of the enclosures and their effects and seldom consider spaceuse. We examined the space use of a group of eight captive chimpanzees 5 years after large-scale enclosure modification at the Johannesburg Zoo, South Africa. Methods: Instantaneous scan sampling was used to record behaviour and location of each chimpanzee at 5 min intervals in the new enclosure. From these 6.8 h of data, space-use patterns and subgroup (two or more chimpanzees within 10 m of each other) spacing were considered relative to local environmental variables, social conditions and the location and size of the previous smaller enclosures in which they had been kept. Results: Space use was heterogeneous, with some enclosure zones being used more than others, and 97.5% of subgroups restricted their spacing to the dimensions of the previous housing (10 m × 10 m). Conclusions: This pattern was not explained by individual behaviour, time of day, location, available space, weather, temperature or shade availability, inter-individual spacing or subgroup composition. We suggest the learned helplessness phenomenon may explain these observations and discuss the implications for both animal welfare and endangered species conservation.Regardless of the mechanism, we suggest that such effects could be avoided through the provision of large enclosures for captive animals.

An Evaluation of Ethograms Measuring Distinct Features of Enrichment Use by Captive Chimpanzees (Pan troglodytes).

Simple SummaryEnvironmental enrichment for chimpanzees is important in order to minimize boredom and stress in captivity and to provide opportunities to engage in species-typical behaviors. However, few studies have investigated potential associations between enrichment objects, manipulation behaviors, and social contexts, nor have they examined if individual chimpanzees vary in their enrichment object preferences. In the current study, three ethograms were used to code the use of enrichment objects, engagement in manipulation behaviors, and social contexts of enrichment use of captive chimpanzees (Pan troglodytes). Data mining from a video archive consisting of 2054 videos collected over a decade allowed the ethograms to be applied to 732 min and 58 s of videos. Some enrichment objects were more often associated with specific manipulation behaviors and social contexts, indicating that enrichment objects might serve distinct social and behavioral purposes. The chimpanzees differed in their enrichment object preferences, suggesting that caregivers of captive chimpanzees should consider individual needs when providing enrichment in order to improve chimpanzees’ experiences in captivity. Finally, the majority of enrichment object use and manipulation behaviors were able to be categorized, indicating that our ethograms were largely effective in coding enrichment use.Environmental enrichment provides mental stimulation and minimizes abnormal behaviors in captive animals. In captive chimpanzees, individual animals may vary in the ways in which they benefit from enrichment or use enrichment devices, so investigating nuances in enrichment use may improve the welfare of captive chimpanzees. In the current study, three ethograms measuring distinct features of enrichment use (i.e., enrichment object, manipulation behavior, and social context) were evaluated by coding videos of captive chimpanzees (Pan troglodytes) at Chimpanzee Sanctuary Northwest in Cle Elum, WA. A total of 732 min and 58 s of video footage was coded from a larger video archive (i.e., 2054 videos) of enrichment use that spanned a decade. A principal component analysis (PCA) revealed that different categories of enrichment objects were more often associated with specific manipulation behaviors and social contexts, suggesting that enrichment objects might fulfill different behavioral and social needs in captivity. Specifically, toy objects were associated with active tactile behaviors in affiliative contexts while oral behaviors were used with foraging objects in solitary contexts. Additionally, individual chimpanzees showed unique preferences for enrichment objects, indicating that caregivers of captive chimpanzees should consider individual needs instead of a “one size fits all” approach to enrichment provisions.

The social dynamics of complex gestural communication in great and lesser apes (Pan troglodytes, Pongo abelii, Symphalangus syndactylus).

Gestures play an essential role in primate communication. However, little is known about how complexity of gestural use (in terms of repertoire size, intentional use, flexibility and use of gestural sequences) relates to individual and dyadic measures of sociality and whether more complex gestural use is more effective in eliciting a response. We observed 19 captive chimpanzees (Pan troglodytes), 16 Sumatran orangutans (Pongo abelii) and 18 siamangs (Symphalangus syndactylus) to assess the complexity and effectiveness of their gestural use. We found that, beyond interspecies variation, the number of gesture types used in a dyad was higher when individuals had stronger social bonds; the probability of accounting for others' attention increased with age, especially for visual gestures; and sequences were more likely used by younger or socially less integrated individuals. In terms of effectiveness, older individuals and those using fewer sequences were more likely to be responded to, while across dyads, the probability of obtaining a response was higher when both individuals accounted for the other's attention and when they used fewer sequences. Overall, this confirms the link between sociality and complex gestural use and suggests that more complex forms of communication, at least in terms of intentional use, may be more effective at achieving communicative goals. This article is part of the theme issue 'Cognition, communication and social bonds in primates'.

Correction for Sharp et al., "Simian Immunodeficiency Virus Infection of Chimpanzees".

Volume 79, no. 7, 3891–3902, 2005, https://doi.org/10.1128/JVI.79.7.3891-3902.2005. Page 3897, Fig. 6B: It has recently been pointed out to us that the Western blot strips depicting the uninfected captive chimpanzee samples Ch-To and C425 appear identical to those from the uninfected wild-living chimpanzee samples Ch-13 and Ch-04. As noted in the figure legend, the data from captive chimpanzees was adapted from Fig. 2A of M. L. Santiago, M. Lukasik, S. Kamenya, Y. Li, et al. (J Virol 77:7545–7562, 2003, https://doi.org/10.1128/jvi.77.13.7545-7562.2003). Inspection of Fig. 2A in Santiago et al. (2003) indicates that the strips for samples Ch-To and C425 do not match those shown in our Fig. 6B. Thus, we conclude that the strips shown for samples Ch-13 and Ch-04 are correct, and these have been inadvertently duplicated in the artwork of Fig. 6B as strips from samples Ch-To and C425; for the correct strips for samples Ch-To and C425, please consult Fig. 2A in Santiago et al. (2003). All four of these samples from uninfected chimpanzees were negative, and this error in the preparation of the figure has no bearing on any of the conclusions reached in our minireview.

Body mass and growth rates in captive chimpanzees (Pan troglodytes) cared for in African wildlife sanctuaries, zoological institutions, and research facilities.

Captive chimpanzees (Pan troglodytes) mature earlier in body mass and have a greater growth rate compared to wild individuals. However, relatively little is known about how growth parameters compare between chimpanzees living in different captive environments. To investigate, body mass was measured in 298 African sanctuary chimpanzeesand was acquired from 1030 zoological and 442 research chimpanzees, using data repositories. An analysis of covariance, adjusting for age, was performed to assess same-sex body mass differences between adult sanctuary, zoological, and research populations. Piecewise linear regression was performed to estimate sex-specific growth rates and the age at maturation, which were compared between sexes and across populations using extra-sum-of-squares F tests. Adult body mass was greater in the zoological and resarch populations compared to the sanctuary chimpanzees, in both sexes. Male and female sanctuary chimpanzees were estimated to have a slower rate of growth compared with their zoological and research counterparts. Additionally, male sanctuary chimpanzees were estimated to have an older age at maturation for body mass compared with zoological and research males, whereas the age at maturation was similar across female populations. For both the zoological and research populations, the estimated growth rate was greater in males compared to females. Together, these data contribute to current understanding of growth and maturation in this species and suggest marked differences between the growth patterns of chimpanzees living in different captive environments.

Olfactory Inspection of Female Reproductive States in Chimpanzees

In mammalian species, olfactory cues are important for within and between species communication. These cues can be part of multimodal signals indicating, for example, female fertility potentially perceived by male conspecifics. However, a large gap exists in our understanding of multimodal signaling in non-human catarrhines. Chimpanzees (Pan troglodytes) live in fission-fusion societies, mate promiscuously and express high levels of male-male competition. Females show a striking visual signal, an exaggerated sexual swelling, known to be a proxy of ovulation, while the maximum swelling is not matching the exact time of ovulation. The question remains if males use additional olfactory cues when being able to approach females closely. This would allow males to pinpoint the exact timing of ovulation and could be one reason why high-ranking males sire offspring more successfully than other males. Here, we present the first systematic test of such multimodal signaling by investigating male sniffing behavior directed to females in relation to their fertility in a group of 13 captive chimpanzees. Our results show that male sniffing behavior significantly increased with female swelling size, with female age as well as when fewer male competitors were present. Hence, odors might be part of a multimodal fertility cue, supporting the idea that males monitor both visual and olfactory cues to gain comprehensive information on female fertility.

Genetic determinants of individual variation in the superior temporal sulcus of chimpanzees (Pan troglodytes).

The superior temporal sulcus (STS) is a conserved fold that divides the middle and superior temporal gyri. In humans, there is considerable variation in the shape, folding pattern, lateralization, and depth of the STS that have been reported to be associated with social cognition and linguistic functions. We examined the role that genetic factors play on individual variation in STS morphology in chimpanzees. The surface area and depth of the STS were quantified in sample of 292 captive chimpanzees comprised of two genetically isolated population of individuals. The chimpanzees had been previously genotyped for AVPR1A and KIAA0319, two genes that play a role in social cognition and communication in humans. Single nucleotide polymorphisms in the KIAA0319 and AVPR1A genes were associated with average depth as well as asymmetries in the STS. By contrast, we found no significant effects of these KIA0319 and AVPR1A polymorphism on surface area and depth measures for the central sulcus. The overall findings indicate that genetic factors account for a small to moderate amount of variation in STS morphology in chimpanzees. These findings are discussed in the context of the role of the STS in social cognition and language in humans and their potential evolutionary origins.

Chimpanzee (Pan troglodytes) gaze is conspicuous at ecologically-relevant distances

Chimpanzee (Pan troglodytes) sclera appear much darker than the white sclera of human eyes, to such a degree that the direction of chimpanzee gaze may be concealed from conspecifics. Recent debate surrounding this topic has produced mixed results, with some evidence suggesting that (1) primate gaze is indeed concealed from their conspecifics, and (2) gaze colouration is among the suite of traits that distinguish uniquely social and cooperative humans from other primates (the cooperative eye hypothesis). Using a visual modelling approach that properly accounts for specific-specific vision, we reexamined this topic to estimate the extent to which chimpanzee eye coloration is discriminable. We photographed the faces of captive chimpanzees and quantified the discriminability of their pupil, iris, sclera, and surrounding skin. We considered biases of cameras, lighting conditions, and commercial photography software along with primate visual acuity, colour sensitivity, and discrimination ability. Our visual modeling of chimpanzee eye coloration suggests that chimpanzee gaze is visible to conspecifics at a range of distances (within approximately 10 m) appropriate for many species-typical behaviours. We also found that chimpanzee gaze is discriminable to the visual system of primates that chimpanzees prey upon, Colobus monkeys. Chimpanzee sclera colour does not effectively conceal gaze, and we discuss this result with regard to the cooperative eye hypothesis, the evolution of primate eye colouration, and methodological best practices for future primate visual ecology research.

Spontaneous nocturnal erections and masturbation in captive male chimpanzees (Pan troglodytes)

Abstract Nocturnal penile tumescence (NPT) has been reported for males of only a few mammalian species, including humans, albeit this scarcity might be an artefact of the difficulty of documenting it. We investigated NPT in 12 adult male chimpanzees living in an all-male group at Kumamoto Sanctuary, Japan. Recorded non-invasively with infrared video cameras across 72 nights (6 per individual), we observed NPT in two thirds of the individuals (8/12), with events in total. More than three quarters occurred during a transition from sleep to wakefulness, suggesting NPT may be associated with rapid eye movement sleep, similar to humans. Masturbation occurred in conjunction with NPT, including the likely consumption of ejaculate, in 4–6 individuals. While detailing nocturnal erections in chimpanzees may help us to better understand this robust physiological phenomenon in humans, further comparative research is necessary to reconstruct its phylogenetic history. At present, our findings support hypothetical functional explanations such as the facilitation of nocturnal emissions in order to increase ejaculate quality or an increase in tissue oxygenation to prevent erectile dysfunction.

What Does a Notion of Weight Mean to Chimpanzees?

How to best understand how animals think about the world? Considering that cognition develops in interactions with the outside world, we need to compare the cognitive performance of individuals of the same species living under different ecological conditions. I propose here to compare the notion of weight studied in captive chimpanzees to the ones observed in a wild chimpanzee population that is using hammers of different weights to crack nuts during many months of the year. Detailing different situations where the chimpanzees have to select a hammer for cracking two species of nuts of different hardness, my colleagues and I could illustrate the flexibility of the causal reasoning performed by the chimpanzees, including stable anticipation of weight, combining weight with density and transport distance, and the gradual shift of selection criteria with increasing transport distance. Combined with results from captive studies on the notion of weight in chimpanzees, this suggests that when individuals face specific ecological challenges, they may acquire sophisticated causal reasoning generalized over different contexts. The evolution of cognition in chimpanzees suggests a complex interplay with ecology, memory demands, individual differences, and technical challenges important to the population.

Chimpanzee Extraversion scores vary with epigenetic modification of dopamine receptor gene D2 (DRD2) and early rearing conditions

ABSTRACT Chimpanzees have consistent individual differences in behaviour, also referred to as personality. Similar to human personality structure, five dimensions are commonly found in chimpanzee studies that show evidence for convergent and predictive validity (Dominance, Openness, Extraversion, Agreeableness, and Reactivity/Undependability). These dimensions are to some extent heritable, indicating a genetic component that explains part of the variation in personality scores, but are also influenced by environmental factors, such as the early social rearing background of the individuals. In this study, we investigated the role of epigenetic modification of the dopamine receptor D2 gene (DRD2) as a potential mechanism underlying personality variation in 51 captive chimpanzees. We used previously collected personality trait rating data and determined levels of DRD2 CpG methylation in peripheral blood samples for these same individuals. Results showed that DRD2 methylation is most strongly associated with Extraversion, and that varying methylation levels at specific DRD2 sites are associated with changes in Extraversion in nursery-reared, but not mother-reared, individuals. These results highlight the role of dopaminergic signalling in chimpanzee personality, and indicate that environmental factors, such as social experiences early in life, can have long-lasting behavioural effects, potentially through modification of the epigenome. These findings add to the growing evidence demonstrating the importance of the experience-dependent methylome for the development of complex social traits like personality.