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Empathy is a complex, multi‐dimensional capacity that facilitates the sharing and understanding of others' emotions. As our closest living relatives, bonobos (Pan paniscus) and chimpanzees (P. troglodytes) provide an opportunity to explore the origins of hominin social cognition, including empathy. Despite certain assumptions that bonobos and chimpanzees may differ empathically, these species appear to overlap considerably in certain socio‐emotional responses related to empathy. However, few studies have systematically tested for species variation in Pan empathic or socio‐emotional tendencies. To address this, we synthesise the growing literature on Pan empathy to inform our understanding of the selection pressures that may underlie the evolution of hominin empathy, and its expression in our last common ancestor. As bonobos and chimpanzees show overlaps in their expression of complex socio‐emotional phenomena such as empathy, we propose that group comparisons may be as or more meaningful than species comparisons when it comes to understanding the evolutionary pressures for such behaviour. Furthermore, key differences, such as how humans and Pan communicate, appear to distinguish how we experience empathy compared to our closest living relatives. [ABSTRACT FROM AUTHOR]

Primate social organizations, or grouping patterns, vary significantly across species. Behavioral strategies that allow for flexibility in grouping patterns offer a means to reduce the costs of group living. Chimpanzees (Pan troglodytes) have a fission‐fusion social system in which temporary subgroups ("parties") change in composition because of local socio‐ecological conditions. Notably, western chimpanzees (P. t. verus) are described as showing a higher degree of bisexual bonding and association than eastern chimpanzees, and eastern female chimpanzees (P. t. schweinfurthii) are thought to be more solitary than western female chimpanzees. However, reported comparisons in sociality currently depend on a small number of study groups, particularly in western chimpanzees, and variation in methods. The inclusion of additional communities and direct comparison using the same methods are essential to assess whether reported subspecies differences in sociality hold in this behaviorally heterogeneous species. We explored whether sociality differs between two communities of chimpanzees using the same motion‐triggered camera technology and definitions of social measures. We compare party size and composition (party type, sex ratio) between the western Gahtoy community in the Nimba Mountains (Guinea) and the eastern Waibira community in the Budongo Forest (Uganda). Once potential competition for resources such as food and mating opportunities were controlled for, subspecies did not substantially influence the number of individuals in a party. We found a higher sex‐ratio, indicating more males in a party, in Waibira; this pattern was driven by a greater likelihood in Gahtoy to be in all‐female parties. This finding is the opposite of what was expected for eastern chimpanzees, where female‐only parties are predicted to be more common. Our results highlight the flexibility in chimpanzee sociality, and caution against subspecies level generalizations. Research highlights: Party size in western and eastern chimpanzees did not differ between our two study communities suggesting that the number of individuals in a party is shaped by similar socio‐ecological pressures.In our sample, eastern chimpanzees were found to have more males in a party compared to western chimpanzees, contrary to the expectation of more female‐only parties in eastern compared to western chimpanzees.Our findings highlight the flexibility in chimpanzee grouping patterns and caution against subspecies level generalizations from limited data points and/or heterogeneous methods. [ABSTRACT FROM AUTHOR]

Compassion fatigue (CF) refers to the exhaustion and distress caused by the demands of caring for others. CF occurs in a variety of helping professions, including physicians, nurses, educators, social workers and animal caregivers, and is known to adversely impact both caregivers' quality of life and the care they provide. This study assessed the prevalence, risk and protective factors, coping strategies and support programs for CF in chimpanzee caregivers (N = 123) at accredited sanctuaries and zoos in the United States. Online survey results revealed that 91.06% of chimpanzee caregivers experienced CF at some point in their careers. Common CF symptoms were exhaustion, frustration, anxiety, depression, and apathy. Perceived factors influencing CF included being understaffed, lacking resources and training, poor relationships with coworkers and supervisors, and financial insecurity. Commonly reported coping strategies were talking to someone, having pets, self-care, and getting away from work. 20.33% of caregivers reported having institutional support programs available to them, however they were rarely viewed as helpful and 32.52% of respondents were unsure about program availability. Overall, our findings suggest that, like other caregiving professionals, chimpanzee caregivers are susceptible to CF and may benefit from new or updated support programs that continue to build a 'culture of care' that meets employee, animal, and facility needs. [ABSTRACT FROM AUTHOR]

Simple Summary: The aim of protected areas is to conserve nature. However, the wild animal populations they shelter are not immune to extinction. It is, therefore, important for researchers to provide regular information on the ecology of these populations in order to guide management actions and enhance knowledge. From studies based on chimpanzee nests, we revealed that chimpanzees in Niokolo Koba National Park nest more in closed habitats than in open ones, and that certain plant species are preferentially chosen for nest construction. Diospyros mespiliformis and Pterocarpus erinaceus are the trees most used by chimpanzees for bearing their nests. In addition, we found that chimpanzee nest height and nest life were influenced by the environmental factors. We conclude that the preservation of nesting sites of chimpanzees is crucial for the conservation of this great ape, considered to be on the border of extinction in Senegal. The Niokolo Koba National Park (NKNP) is the largest protected area in Senegal and lies at the northern limit of the chimpanzee's range in West Africa. Recent information on nesting behavior and factors influencing nesting behavior is available for several sites outside NKNP. However, the information available for NKNP is obsolete. Considering that the adequate management of chimpanzee populations cannot be achieved without strong scientific knowledge, it is essential to update data on chimpanzee, Pan troglodytes verus, nesting behavior in NKNP. For this reason, we surveyed their habitat in Mt. Assirik and recorded 626 chimpanzee nests. The results of the study showed that chimpanzees nest more often in closed-canopy habitats such as gallery forests. The average nest height observed in this study was 8.07 ± 0.36 m, varying between 2 and 20 m, which is well below the heights described in most sites where chimpanzees cohabit with large carnivores. Botanical surveys confirmed that chimpanzees select tree species bearing their nests. In Assirik, 12 of the 37 tree species bearing nests are the most used. The nest decay rate (or the time it takes for a nest to go from the fresh to the skeletal stage) at Assirik averaged 68.8 ± 5.8 days. [ABSTRACT FROM AUTHOR]

We report the presence of habitual ground nesting in a newly studied East African chimpanzee (Pan troglodytes schweinfurthii) population in the Bugoma Central Forest Reserve, Uganda. Across a 2‐year period, we encountered 891 night nests, 189 of which were classified as ground nests, a rate of ~21%. We find no preliminary evidence of socio‐ecological factors that would promote its use and highlight local factors, such as high incidence of forest disturbance due to poaching and logging, which appear to make its use disadvantageous. While further study is required to establish whether this behavior meets the strict criteria for nonhuman animal culture, we support the argument that the wider use of population and group‐specific behavioral repertoires in flagship species, such as chimpanzees, offers a tool to promote the urgent conservation action needed to protect threatened ecosystems, including the Bugoma forest. Research highlights: Chimpanzees in the Bugoma Forest, Uganda, habitually employ overnight ground nests (21% of total nests, ground nests present in 25% of nest sites).We discuss a range of socio‐ecological and cultural explanations, highlighting local factors that appear to be disadvantageous for the behavior.While behavioral variation may offer a powerful tool for conservation, strict scientific thresholds for establishing cultural explanations may not be compatible with urgent conservation timelines. [ABSTRACT FROM AUTHOR]

Background: We tackle the problem of estimating species TMRCAs (Time to Most Recent Common Ancestor), given a genome sequence from each species and a large known phylogenetic tree with a known structure (typically from one of the species). The number of transitions at each site from the first sequence to the other is assumed to be Poisson distributed, and only the parity of the number of transitions is observed. The detailed phylogenetic tree contains information about the transition rates in each site. We use this formulation to develop and analyze multiple estimators of the species' TMRCA. To test our methods, we use mtDNA substitution statistics from the well-established Phylotree as a baseline for data simulation such that the substitution rate per site mimics the real-world observed rates. Results: We evaluate our methods using simulated data and compare them to the Bayesian optimizing software BEAST2, showing that our proposed estimators are accurate for a wide range of TMRCAs and significantly outperform BEAST2. We then apply the proposed estimators on Neanderthal, Denisovan, and Chimpanzee mtDNA genomes to better estimate their TMRCA with modern humans and find that their TMRCA is substantially later, compared to values cited recently in the literature. Conclusions: Our methods utilize the transition statistics from the entire known human mtDNA phylogenetic tree (Phylotree), eliminating the requirement to reconstruct a tree encompassing the specific sequences of interest. Moreover, they demonstrate notable improvement in both running speed and accuracy compared to BEAST2, particularly for earlier TMRCAs like the human-Chimpanzee split. Our results date the human – Neanderthal TMRCA to be ∼ 408 , 000 years ago, considerably later than values cited in other recent studies. [ABSTRACT FROM AUTHOR]

The outcome of an action often occurs after a delay. One solution for learning appropriate actions from delayed outcomes is to rely on a chain of state transitions. Another solution, which does not rest on state transitions, is to use an eligibility trace (ET) that directly bridges a current outcome and multiple past actions via transient memories. Previous studies revealed that humans (Homo sapiens) learned appropriate actions in a behavioral task in which solutions based on the ET were effective but transition-based solutions were ineffective. This suggests that ET may be used in human learning systems. However, no studies have examined nonhuman animals with an equivalent behavioral task. We designed a task for nonhuman animals following a previous human study. In each trial, participants chose one of two stimuli that were randomly selected from three stimulus types: a stimulus associated with a food reward delivered immediately, a stimulus associated with a reward delivered after a few trials, and a stimulus associated with no reward. The presented stimuli did not vary according to the participants' choices. To maximize the total reward, participants had to learn the value of the stimulus associated with a delayed reward. Five chimpanzees (Pan troglodytes) performed the task using a touchscreen. Two chimpanzees were able to learn successfully, indicating that learning mechanisms that do not depend on state transitions were involved in the learning processes. The current study extends previous ET research by proposing a behavioral task and providing empirical data from chimpanzees. [ABSTRACT FROM AUTHOR]

Honest signalling theory suggests that humans and chimpanzees can extract socially relevant information relating to personality from the faces of their conspecifics. Humans are also able to extract information from chimpanzees' faces. Here, we examine whether personality characteristics of chimpanzees, including measures of psychopathy, can be discerned based purely on facial morphology in photographs. Twenty-one chimpanzees were given naïve and expert personality ratings on the Ten Item Personality Inventory (TIPI) and the Chimpanzee Triarchic Model of Psychopathy (CHMP-Tri) before and following behavioural observations. Characteristics relating to openness, conscientiousness, extraversion, and disinhibition could be distinguished from the faces of chimpanzees. Individuals higher on disinhibition have lower scores on conscientiousness and emotional stability and higher scores on extraversion, while those higher on meanness have lower conscientiousness and agreeableness. Facial expressions are linked to personality traits present in the TIPI and CHMP-Tri models: the Relaxed Face and the Grooming Face were displayed more by chimpanzees higher on agreeableness, whereas the Compressed Lips Face was observed more in those individuals higher on boldness, and the Full Open Grin was displayed more by chimpanzees higher on extraversion but lower on emotional stability and conscientiousness. Facial expressions were also found to be associated with particular behavioural contexts, namely the Grooming Face in affiliative contexts and the Relaxed and Relaxed Open Mouth Faces in neutral contexts. Dominant chimpanzees display higher levels of boldness and more Compressed Lips Faces, Relaxed Open Mouth Faces, and Grooming Faces than subordinate individuals. These findings support and extend evidence for an honest signalling system and a personality structure shared between humans and chimpanzees. Future research could further explore how personality is conveyed through the face, perhaps through more than just singular aspects of character, and maybe reflecting what chimpanzees themselves are able to do. [ABSTRACT FROM AUTHOR]

Simple Summary: Chimpanzees Tatu and Loulis use signs of American Sign Language. This study presents sign use over 8 years while living at the sanctuary of the Fauna Foundation. They used a majority of the signs in their base vocabulary each year in a variety of communicative functions. Adult chimpanzees Tatu and Loulis lived at the Fauna Foundation sanctuary. They had acquired signs of American Sign Language (ASL) while young and continued to use them as adults. Caregivers with proficiency in ASL maintained daily sign language records during interactions and passive observation. Sign checklists were records of daily vocabulary use. Sign logs were records of signed interactions with caregivers and other chimpanzees. This study reports sign use from eight years of these records. Tatu and Loulis used a majority of their base vocabularies consistently over the study period. They used signs that they had acquired decades earlier and new signs. Their utterances served a variety of communicative functions, including responses, conversational devices, requests, and descriptions. They signed to caregivers, other chimpanzees, including those who did not use signs, and to themselves privately. This indicates the importance of a stimulating and interactive environment to understand the scope of ape communication and, in particular, their use of sign language. [ABSTRACT FROM AUTHOR]

This article discusses the detection of natural non-polio enteroviruses (EVs) in captive mandrills and chimpanzees housed in a primate center in Franceville, Gabon. The study found that EVs were present in both species, with mandrills carrying EVs of the species EV-J and chimpanzees carrying EVs of the species EV-A90. The detection of EVs in mandrills did not pose a threat to the species, as healthy captive and wild individuals also harbored these EVs. However, the detection of EV-A90 in chimpanzees highlighted the potential risk of zoonotic transmission from humans. The study emphasizes the importance of monitoring and studying EVs in order to protect primate health and prevent the introduction of human pathogens into the wild. [Extracted from the article]

Objectives: To understand the function of food sharing among our early hominin ancestors, we can turn to our nonhuman primate relatives for insight. Here, we examined the function of meat sharing by Fongoli chimpanzees, a community of western chimpanzees (Pan troglodytes verus) in southeastern Sénégal. Materials and Methods: We tested three non‐mutually exclusive hypotheses that have been used to explain patterns of food sharing: kin selection, generalized reciprocity, and meat‐for‐mating opportunities. We analyzed meat sharing events (n = 484) resulting from hunts, along with data on copulations, age‐sex class, and kinship to determine which variables predict the likelihood of meat sharing during this study period (2006–2019). Results: We found full or partial support for kin selection, direct reciprocity, and meat‐for‐mating‐opportunities. However, the analyses reveal that reciprocity and a mother/offspring relationship were the strongest predictors of whether or not an individual shared meat. Conclusions: The results of this study emphasize the complexity of chimpanzee meat sharing behaviors, especially at a site where social tolerance offers increased opportunities for meat sharing by individuals other than dominant males. These findings can be placed in a referential model to inform hypotheses about the sensitivity of food sharing to environmental pressures, such as resource scarcity in savanna landscapes. [ABSTRACT FROM AUTHOR]

Testosterone promotes mating effort, which involves intraspecific aggression for males of many species. Therefore, males with higher testosterone levels are often thought to be more aggressive. For mammals living in multimale groups, aggression is hypothesized to link male social status (i.e. dominance rank) and testosterone levels, given that high status predicts mating success and is acquired partly through aggressive intragroup competition. In male chimpanzees, Pan troglodytes , dominance rank has been repeatedly linked to interindividual variation in testosterone levels, but evidence directly linking interindividual variation in testosterone and aggression is lacking. In the present study, we test both aggression levels and lean muscle mass, as measured by urinary creatinine, as links between dominance rank and testosterone levels in a large sample of wild male chimpanzees. Multivariate analyses indicated that dominance rank was positively associated with total rates of intragroup aggression, average urinary testosterone levels and average urinary creatinine levels. Testosterone was positively associated with creatinine levels but negatively associated with total aggression rates. Furthermore, mediation analyses showed that testosterone levels facilitated an association between dominance rank and creatinine levels. Our results indicate that (1) adult male chimpanzees with higher average testosterone levels are often higher ranking but not more aggressive than males with lower testosterone and (2) lean muscle mass links dominance rank and testosterone levels in Ngogo males. We assert that aggression rates are insufficient to explain links between dominance rank and testosterone levels in male chimpanzees and that other social variables (e.g. male–male relationship quality) may regulate testosterone's links to aggression. • We examined rank, testosterone, aggression and creatinine in male chimpanzees. • Urinary testosterone levels were positively associated with male dominance rank. • Urinary testosterone levels were negatively associated with aggression rates. • Urinary creatinine was positively associated with dominance rank and testosterone. • Urinary testosterone mediated dominance rank's association with urinary creatinine. [ABSTRACT FROM AUTHOR]

Diagnosis and treatment of atopic dermatitis (AD) in chimpanzees are challenging. Validated allergy tests specific for chimpanzees are not available. A multifactorial management of atopic dermatitis is important. Successful management of AD has, to the best knowledge of the authors, not been described in chimpanzees. [ABSTRACT FROM AUTHOR]

Directedness and engagement during pre‐verbal vocal communication play a major role in language development. What was their role in the evolution of language? This question invites us to examine these behaviours in chimpanzee vocal ontogeny. We collected observational data on infant (N = 15) and juvenile (N = 13) chimpanzees at Chimfunshi Wildlife Orphanage, Zambia. We examined the impact of age and vocalization type (grunts, whimpers, laughs and screams) on directed cues (gaze directedness and face directedness) and engagement (mutual face directedness) during vocal communication. We also assessed the impact of directed cues and engagement on social interactions by coding the behaviour of social partners before, during and after a vocalisation, and examining whether they contingently changed their behaviour in response to the vocalisation if it was directed or if engagement occurred. We found that face directed vocalisations showed a general increase during ontogeny and we observed call‐type dependent effects of age for mutual face directedness. Only face directed vocalisations were significantly predictive of behavioural responses in social partners. We conclude that like young humans, young chimpanzees routinely exhibit directed behaviours and engagement during vocal communication. This social competency improves during ontogeny and benefits individuals by increasing the chances of eliciting behavioural responses from social partners. Directedness and engagement likely provide a foundation for language phylogenetically, as well as ontogenetically. Research Highlights: We show that directedness and engagement routinely occur during early chimpanzee vocalisations.Directedness increases throughout chimpanzee vocal ontogeny, similar to human infants.Directedness enhances social partner responsiveness, demonstrating a direct benefit to this style of communication.Directedness and engagement could provide a route towards language phylogenetically as well as ontogenetically. [ABSTRACT FROM AUTHOR]

A 41‐year‐old male vasectomized, zoo‐housed chimpanzee (Pan troglodytes) presented with progressive visual deficits due to bilateral cataract formation. Phacoemulsification and lenticular implant were performed by a veterinary and human board‐certified ophthalmologist team in a field setting. Post‐operative healing occurred without complication, and the patient returned to the troop with improved vision. [ABSTRACT FROM AUTHOR]

Effective species conservation management relies on detailed species distribution data. For many species, such as chimpanzees (Pan troglodytes), distribution data are collected during ground surveys. For chimpanzees, such ground surveys usually focus on detection of the nests they build instead of detection of the chimpanzees themselves due to their low density. However, due to the large areas they still occur in, such surveys are very costly to conduct and repeat frequently to monitor populations over time. Species distribution models are more accurate if they include presence as well as absence data. Earlier studies used drones to determine chimpanzee presence using nests. In this study, therefore, we explored the use of drones to determine the absence of chimpanzee nests in areas we flew over on the edge of the chimpanzee distribution in western Tanzania. We conducted 13 flights with a fixed-wing drone and collected 3560 images for which manual inspection took 180 h. Flights were divided into a total of 746 25 m2 plots for which we determined the absence probability of nests. In three flights, we detected nests, in eight, absence was assumed based on a 95% probability criterion, and in two flights, nest absence could not be assumed. Our study indicates that drones can be used to cover relatively large areas to determine the absence of chimpanzees. To fully benefit from the usage of drones to determine the presence and absence of chimpanzees, it is crucial that methods are developed to automate nest detection in images. [ABSTRACT FROM AUTHOR]

Male and female human social bonding strategies are culturally shaped, in addition to being genetically rooted. Investigating nonhuman primate bonding strategies across sex groups allows researchers to assess whether, as with humans, they are shaped by the social environment or whether they are genetically predisposed. Studies of wild chimpanzees show that in some communities males have strong bonds with other males, whereas in others, females form particularly strong intrasex bonds, potentially indicative of cultural differences across populations. However, excluding genetic or ecological explanations when comparing different wild populations is difficult. Here, we applied social network analysis to examine male and female social bonds in two neighbouring semiwild chimpanzee groups of comparable ecological conditions and subspecies compositions, but that differ in demographic makeup. Results showed differences in bonding strategies across the two groups. While female–female party co-residence patterns were significantly stronger in Group 1 (which had an even distribution of males and females) than in Group 2 (which had a higher proportion of females than males), there were no such differences for male–male or male–female associations. Conversely, female–female grooming bonds were stronger in Group 2 than in Group 1. We also found that, in line with captive studies but contrasting research with wild chimpanzees, maternal kinship strongly predicted proximity and grooming patterns across the groups. Our findings suggest that, as with humans, male and female chimpanzee social bonds are influenced by the specific social group they live in, rather than predisposed sex-based bonding strategies. [ABSTRACT FROM AUTHOR]

Simple Summary: This study aimed to teach the numerical sequence from 1 to 19 in the decimal system to six chimpanzees. The participants were three mother–child pairs. The original goal for chimpanzees was to touch the numerals on the display from 1 to 19, in this order. The baseline daily training was twofold: touching the adjacent numerals from 1 to X and X to 19 in ascending order. In two separate ways, chimpanzees succeeded at touching adjacent numerals in the range 1 to 19. Systematic tests assessed four factors: range (1 to 9 vs. 1 to 19), adjacency (adjacent vs. nonadjacent numerals), number of stimuli used (three, four, and five), and memory load (nonmemory vs. a memory task called the "masking task"). All four factors were important. A further test directly compared the performance of chimpanzees with that of human participants using the same apparatus and procedure. Both accuracy and response latency showed that processing two-digit numerals was more difficult than one-digit numerals in both species. A chimpanzee named Pal perfectly mastered the order of two-digit numerals just like humans. The difference between the two species was discussed in terms of species-specific global–local information processing. The sequence of Arabic numerals from 1 to 19 was taught to six chimpanzees, three pairs of mother and child. Each chimpanzee participant sat facing a touchscreen on which the numerals appeared in random positions within an imaginary 5-by-8 matrix. They had to touch the numerals in ascending order. Baseline training involved touching the adjacent numerals from 1 to X or from the numeral X to 19. Systematic tests revealed the following results: (1) The range 1 to 9 was easier than 1 to 19. (2) Adjacent numerals were easier than nonadjacent ones. (3) The "masking" (memory task) caused deterioration of performance. All these factors depended on the number of numerals simultaneously presented on the screen. A chimpanzee named Pal mastered the skill of ordering two-digit numerals with 100% accuracy. Human participants were tested in the same experiment with the same procedure. Both species showed relative difficulty in handling two-digit numerals. Global–local information processing is known to be different between humans and other primates. The assessment of chimpanzee performance and comparison with humans were discussed in terms of the possible difference in the global–local dual information processing of two-digit numerals. [ABSTRACT FROM AUTHOR]

Simple Summary: Ensuring proper sleep is essential for the health of all animals, including chimpanzees. Many studies have demonstrated that sleep plays a crucial role in physical and behavioral regulation, and sleep disruptions may cause negative consequences for both. This study explores how environmental conditions and nocturnal disturbances affect the sleep and daytime behavior of sanctuary-housed chimpanzees, respectively. Our results indicate that indoor temperature and humidity affect chimpanzee nocturnal activity. Furthermore, the frequency and intensity of nocturnal disturbances influence the chimpanzee's behavior the following day, i.e., time spent in inactivity, abnormal, self-directed, and affiliative behaviors. These findings highlight the importance of controlling factors that affect the quality of chimpanzees' sleep at night. The use of affordable audio recordings has the potential to provide valuable information regarding the chimpanzee's nocturnal activity and wellbeing. Chimpanzee's nocturnal sleep is a dynamic and complex process, still not fully understood. As in humans, not getting enough quality sleep due to frequent or lasting disruptions may affect their physical and mental health, hence wellbeing, which may be reflected in their daytime behavior. This study aims to understand the impact of abiotic factors, such as temperature and humidity on the nocturnal activity as well as the impact of nocturnal awakening events on daytime behavior in sanctuary-housed chimpanzees. We monitored noisy nocturnal activity through audio recordings for one year, documenting the number and duration of sound events produced by chimpanzees to indicate sleep fragmentation and disruption intensity, respectively. Our results indicate that indoor temperature and humidity indeed influence the chimpanzee's nocturnal activity. Furthermore, sleep fragmentation and intensity of nocturnal events significantly influenced the following day's behavior. After nights marked by frequent and/or intense sleep disruptions, higher levels of inactivity, and abnormal and self-directed behaviors were observed, and chimpanzees spent more time on affiliative interactions and in social proximity. These findings highlight the importance of controlling factors influencing nocturnal sleep quality. Furthermore, we demonstrated that economic audio recordings used to monitor nocturnal activity, provide insights into the chimpanzee's behavior and wellbeing. [ABSTRACT FROM AUTHOR]

Simple Summary: Wild chimpanzees flexibly adapt their behavior based on many social and environmental aspects of their lives. These include seasonality and food availability as well as aspects regarding their communities and parties, such as group size, sex ratio and the presence of sexually receptive females. This results in a stimulating but also stressful life. On the contrary, housing conditions in captivity are far less stimulating, often lacking the enriching social component of fluctuating group compositions found in wild fission–fusion societies. Providing captive chimpanzees with an adequate social environment is crucial to ensure their wellbeing. We had the rare opportunity to analyze behavioral data of an all-male chimpanzee group that experienced two major group alterations, i.e., the integration of two adult females and the passing away of two adult males. Our findings highlight the importance of conducting longitudinal observations to objectively record variations in the chimpanzees' behavior. Furthermore, we were able to demonstrate the impact of their social environment, i.e., group size and sex ratio, on the chimpanzees' activity levels and occurrence of abnormal and self-directed behaviors. 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. [ABSTRACT FROM AUTHOR]

Simple Summary: A survey was sent to zoos, research facilities, and sanctuaries which housed chimpanzees. The behavioral profiles of 1122 chimpanzees were collected for this survey. Data collected included information about the animals' age, sex, social group size, rearing history, and enclosure as well as information about each animal's behavior. Each respondent was asked to indicate if certain behaviors had been observed in each chimpanzee over the prior two years. Species typical behaviors (STBs) were queried, including copulation, tool-use, nest-building, and social grooming. Tool-use was reported to be present for 94.3% of the sample, active grooming for 85.7%, copulation for 68.3% and nest-building for 58.9%. Male chimpanzees who were not reared by their conspecific mother were most likely to have deficits in STBs, and female chimpanzees who were mother-reared were generally the most likely to engage in STBs. A survey was sent to zoos, research facilities, and sanctuaries which housed chimpanzees. Data collected included information about 1122 chimpanzees' age, sex, social group-size, rearing history, and enclosure. Respondents were also asked to indicate if certain behaviors had been observed in each chimpanzee over the prior two years. Species- typical behaviors (STBs) were queried, including copulation, tool-use, nest-building, and social grooming. Tool-use was reported present for 94.3% of the sample (n = 982), active social grooming for 85.7% (n = 1121), copulation for 68.3% (n = 863) and nest-building for 58.9% (n = 982). Of the subjects for whom we had data regarding all four STBs (n = 860), 45.6% were reported to engage in all four. Logistic regression analyses using forward Wald criteria were conducted to determine the best model for each STB based on the predictors of age, sex, rearing history, group-size, facility-type, and a sex-by-rearing interaction. The best model for copulation (χ2(6) = 124.62, p < 0.001) included rearing, group-size, facility-type, and the sex-by-rearing interaction. Chimpanzees were more likely to copulate if they were mother-reared, in larger groups, living in research facilities, and, if not mother-reared (NOTMR), more likely to copulate if they were female. The best model for tool-use retained the predictors of age category, facility-type, and sex-by-rearing (χ2(5) = 55.78, p < 0.001). Chimpanzees were more likely to use tools if they were adult, living in research facilities, and if NOTMR, were female. The best model for nest-building included facility-type and rearing (χ2(3) = 205.71, p < 0.001). Chimpanzees were more likely to build nests if they were MR and if they were living in zoos or in sanctuaries. The best model for active social grooming retained the predictors of age, sex, rearing, and type of facility (χ2(6) = 102.15, p < 0.001). Chimpanzees were more likely to engage in active social grooming if they were immature, female, mother-reared, and living in zoos. This research provides a basic behavioral profile for many chimpanzees living under human care in the United States and allows us to determine potential methods for improving the welfare of these and future chimpanzees in this population. [ABSTRACT FROM AUTHOR]

The parietal lobe is a region of especially pronounced change in human brain evolution. Based on comparative neuroanatomical studies, the inferior parietal lobe (IPL) has been shown to be disproportionately larger in humans relative to chimpanzees and macaques. However, it remains unclear whether the underlying histological architecture of IPL cortical areas displays human-specific organization. Chimpanzees are among the closest living relatives of humans, making them an ideal comparative species to investigate potential evolutionary changes in the IPL. We parcellated the chimpanzee IPL using cytoarchitecture and myeloarchitecture, in combination with quantitative comparison of cellular features between the identified cortical areas. Four major areas on the lateral convexity of the chimpanzee IPL (PF, PFG, PG, OPT) and two opercular areas (PFOP, PGOP) were identified, similar to what has been observed in macaques. Analysis of the quantitative profiles of cytoarchitecture showed that cell profile density was significantly different in a combination of layers III, IV, and V between bordering cortical areas, and that the density profiles of these six areas supports their classification as distinct. The similarity to macaque IPL cytoarchitecture suggests that chimpanzees share homologous IPL areas. In comparison, human rostral IPL is reported to differ in its anatomical organization and to contain additional subdivisions, such as areas PFt and PFm. These changes in human brain evolution might have been important as tool making capacities became more complex. [ABSTRACT FROM AUTHOR]

Objectives: This study investigates patterns of bone functional adaptations in extant apes through comparing hindlimb to forelimb bone rigidity ratios in groups with varying levels of arboreality. Materials and Methods: Using CT scans, bone rigidity (J) was calculated at three regions of interest (ROI) along femoral and humeral diaphyses in Homo, Pongo, Pan, and Gorilla with further comparisons made between species and subspecies divisions within Pan and Gorilla. Results: Consistent with previous work on extant hominoids, species exhibited differences in midshaft femoral to humeral (F/H) rigidity ratios. Results of the present study confirm that these midshaft differences extend to 35% and 65% diaphyseal ROIs. Modern humans, exhibiting larger ratios, and orangutans, exhibiting smaller ratios, bracketed the intermediate African apes in comparisons. Within some African apes, limb rigidity ratios varied significantly between taxonomic groups. Eastern gorillas exhibited the highest mean ratios and chimpanzees the lowest at all three ROIs. In posthoc comparisons, chimpanzees and bonobos did not differ in relative limb rigidity ratios at any of the three ROIs. However, western gorillas were more similar to bonobos than eastern gorillas at 50% and 35% ROIs, but not at the 65% ROI. Conclusion: Species, and to a lesser extent subspecies, can be distinguished by F/H limb rigidity ratios according to broad positional behavior patterns at multiple regions of interest along the diaphyses. Similarity of bonobos and western gorillas is in line with behavioral data of bonobos being the most terrestrial of Pan species, and western gorillas the most arboreal of the Gorilla groups. [ABSTRACT FROM AUTHOR]

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 N = 46 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. 夜間陰茎勃起はヒトの男性において広く一般に生じる現象であるが、ヒト以外の動物での報告 は少ない。これは、夜間の観察が難しいことによるものと考えられる。本研究では、京都大学熊 本サンクチュアリにおいて12個体の男性チンパンジーの夜間の様子を赤外線カメラによって 撮影し、各個体につき6夜間分の行動の記録と分析をおこなった。その結果、12個体のうち8 個体で、のべ23夜間において46事例の夜間陰茎勃起が観察された。この46事例のうち11事例 ではマスターベーションを伴って観察された。また、全事例のうち約80％が睡眠から覚醒する過 程において生じており、ヒトと同様にチンパンジーでもレム睡眠の期間中もしくはその後に生じる 現象であることが示唆される。夜間陰茎勃起のあとにマスターベーションをおこなったのは4－6 個体だった。マスターベーションのあとに手を口に運ぶ動作が確認されており、精液を食べた と推測される。チンパンジーの夜間勃起現象の詳細な記述により、ヒトにおける本現象につい ての理解が深まると期待できるが、系統的基盤を明らかにするためには今後さらなる比較研 究が必要である。我々の観察は、夜間勃起現象の機能として、射精を促し古い精子を排出し て新しい精子に入れ替える効果や、組織への酸素供給の効果があるとする仮説を支持するも のである。 [ABSTRACT FROM AUTHOR]

Humans are argued to be unique in their ability and motivation to share attention with others about external entities—sharing attention for sharing’s sake. Indeed, in humans, using referential gestures declaratively to direct the attention of others toward external objects and events emerges in the first year of life. In contrast, wild great apes seldom use referential gestures, and when they do, it seems to be exclusively for imperative purposes. This apparent species difference has fueled the argument that the motivation and ability to share attention with others is a human-specific trait with important downstream consequences for the evolution of our complex cognition [M. Tomasello, Becoming Human (2019)]. Here, we report evidence of a wild ape showing a conspecific an item of interest. We provide video evidence of an adult female chimpanzee, Fiona, showing a leaf to her mother, Sutherland, in the context of leaf grooming in Kibale Forest, Uganda. We use a dataset of 84 similar leaf-grooming events to explore alternative explanations for the behavior, including food sharing and initiating dyadic grooming or playing. Our observations suggest that in highly specific social conditions, wild chimpanzees, like humans, may use referential showing gestures to direct others’ attention to objects simply for the sake of sharing. The difference between humans and our closest living relatives in this regard may be quantitative rather than qualitative, with ramifications for our understanding of the evolution of human social cognition. [ABSTRACT FROM AUTHOR]

Simple Summary: Chimpanzees and humans are both species of hominoid primates that are closely related phylogenetically. One of the key differences between these two species is their use of their upper extremities. Humans use this limb mainly in manipulative tasks, while chimpanzees also use it during locomotion. In this study, we have analyzed the muscle architecture and the expression of the myosin heavy chain isoforms in the two elbow extensor muscles, the triceps brachii and the anconeus, in humans and chimpanzees, in order to find differences that could be related to the different uses of the upper extremities in these species. We have found that the triceps brachii of chimpanzees is more prepared for strength and power as an adaptation to locomotion, while the same muscle in humans is more prepared for speed and resistance to fatigue as an adaptation to manipulative activities. Our results increase the knowledge we have of the musculoskeletal system of chimpanzees and can be applied in various fields, such as comparative anatomy, evolutionary anatomy or anthropology. The anatomical and functional characteristics of the elbow extensor muscles (triceps brachii and anconeus) have not been widely studied in non-human hominoid primates, despite their great functional importance. In the present study, we have analyzed the muscle architecture and the expression of the myosin heavy chain (MHC) isoforms in the elbow extensors in humans and chimpanzees. Our main objective was to identify differences in these muscles that could be related to the different uses of the upper extremity in the two species. In five humans and five chimpanzees, we have analyzed muscle mass (MM), muscle fascicle length (MFL), and the physiological cross-sectional area (PCSA). In addition, we have assessed the expression of the MHC isoforms by RT-PCR. We have found high MM and PCSA values and higher expression of the MHC-IIx isoform in the triceps brachii of chimpanzees, while in humans, the triceps brachii has high MFL values and a higher expression of the MHC-I and MHC-IIa isoforms. In contrast, there were no significant differences between humans and chimpanzees in any of the values for the anconeus. These findings could be related to the participation of the triceps brachii in the locomotion of chimpanzees and to the use of the upper extremity in manipulative functions in humans. The results obtained in the anconeus support its primary function as a stabilizer of the elbow joint in the two species. [ABSTRACT FROM AUTHOR]

Objectives: Energetics are widely recognized to influence timing of birth in humans and other eutherian mammals, yet considerable variation exists in the relationship between energetic constraints and gestation length. In humans, poor nutrition and short inter‐gestational intervals (IGIs) are associated with shorter gestations. In other mammals, lower energy availability is usually associated with longer gestations. We investigated the predictors of gestation length, and the impact of gestation length on offspring survival, in chimpanzees, humans' closest living relatives. Materials and Methods: We used 50 years of demographic and behavioral data to estimate gestation lengths in the wild chimpanzees of Gombe National Park, Tanzania, and then used ecological and demographic data to explore the predictors and consequences of gestation length in our sample. Results: Gestation lengths were shorter for females in their early 30s (relative to younger and older females), and after short IGIs. Other predictors potentially associated with maternal energetic condition and maternal investment were not associated with gestation length. We also found that shorter gestation lengths corresponded to lower offspring survival. Discussion: Like humans, chimpanzees had shorter gestations after short IGIs, and short gestations were associated with higher offspring mortality. We consider competing explanations for the conflicting relationships between energetics and gestation length across eutherian mammals in light of these results. [ABSTRACT FROM AUTHOR]

Vocal learning, the ability to modify the acoustic structure of vocalizations based on social experience, is a fundamental feature of speech in humans (Homo sapiens). While vocal learning is common in taxa such as songbirds and whales, the vocal learning capacities of nonhuman primates appear more limited. Intriguingly, evidence for vocal learning has been reported in chimpanzees (Pan troglodytes), for example, in the form of regional variation ("dialects") in the "pant‐hoot" calls. This suggests that some capacity for vocal learning may be an ancient feature of the Pan‐Homo clade. Nonetheless, reported differences have been subtle, with intercommunity variation representing only a small portion of the total acoustic variation. To gain further insights into the extent of regional variation in chimpanzee vocalizations, we performed an analysis of pant‐hoots from chimpanzees in the neighboring Kasekela and Mitumba communities at Gombe National Park, Tanzania, and the geographically distant Kanyawara community at Kibale National Park, Uganda. We did not find any statistically significant differences between the neighboring communities at Gombe or among geographically distant communities. Furthermore, we found differences among individuals in all communities. Hence, the variation in chimpanzee pant‐hoots reflected individual differences, rather than group differences. Thus, we did not find evidence of dialects in this population, suggesting that extensive vocal learning emerged only after the lineages of Homo and Pan diverged. Highlights: Previous studies have reported dialects in chimpanzee pant‐hoot calls.Vocal dialects may reflect vocal learning ability.We compared two neighboring and one geographically distant chimpanzee communities.The acoustic structure of pant‐hoots reflected individual but not group identity. [ABSTRACT FROM AUTHOR]

For energetically limited organisms, life‐history theory predicts trade‐offs between reproductive effort and somatic maintenance. This is especially true of female mammals, for whom reproduction presents multifarious energetic and physiological demands.Here, we examine longitudinal changes in the gut virome (viral community) with respect to reproductive status in wild mature female chimpanzees Pan troglodytes schweinfurthii from two communities, Kanyawara and Ngogo, in Kibale National Park, Uganda.We used metagenomic methods to characterize viromes of individual chimpanzees while they were cycling, pregnant and lactating.Females from Kanyawara, whose territory abuts the park's boundary, had higher viral richness and loads (relative quantity of viral sequences) than females from Ngogo, whose territory is more energetically rich and located farther from large human settlements. Viral richness (total number of distinct viruses per sample) was higher when females were lactating than when cycling or pregnant. In pregnant females, viral richness increased with estimated day of gestation. Richness did not vary with age, in contrast to prior research showing increased viral abundance in older males from these same communities.Our results provide evidence of short‐term physiological trade‐offs between reproduction and infection, which are often hypothesized to constrain health in long‐lived species. [ABSTRACT FROM AUTHOR]

Environmental enrichment is essential for the well‐being of zoo animals. Recent advances in sensor and video technologies may contribute to improvements in enrichment in terms of their flexibilities and time constraints. The purpose of this study was to investigate whether interactive movie art can be used as a form of environmental enrichment. We implemented interactive movies designed by a professional artist, a visual art aiming to reflect naturalistic forest habitat, in an indoor chimpanzee enclosure at Kyoto City Zoo in Japan. Motion‐tracking sensors embedded in buoys were installed at several locations around the indoor enclosure; the chimpanzees could change the movie contents by physically interacting with these objects. We recorded behaviors by observing entire troop of chimpanzees (six) between March 16 and 20, 2020 (control condition), then recorded behaviors when the interactive movie was presented (experimental condition) between March 21 and 29, 2020. Behaviors were recorded via direct observations and video recordings to examine any changes after the installation of interactive art. The chimpanzees spent more time in the indoor enclosures during the experimental condition than during the control condition. Activity budgets did not change substantially during the study period. There was no evidence of habituation to the movie during the study period. Three chimpanzees, including two young chimpanzees, interacted with the movie more frequently than the others; these young chimpanzees occasionally showed playful expressions when interacting with the movie and exhibited different reactivities to the movie scenes. These results demonstrate, first, that the interactive art did not negatively affect chimpanzee behavior, and second, that some of the chimpanzees indeed showed positive responses to the art. This study, therefore, introduces a novel possibility for environmental enrichment in zoos, involving a collaboration between science and art. Research Highlights: We investigate whether technology‐based interactive movie art can be used as a form of environmental enrichment by analyzing their behaviors.The chimpanzees spent more time in the locations where the arts were presented and did not lose interest across the experimental condition. Two young chimpanzees particularly used it and showed play face in relation to the interactive arts.These results demonstrate that the interactive art did not negatively affect chimpanzee behavior, and that some of the chimpanzees indeed showed positive responses to the art. [ABSTRACT FROM AUTHOR]

Examining the links between intentional communication and social relationships provides insights into the cognitive skills needed to manage a differentiated set of social bonds. Great apes gesture intentionally, but how this intentionality relates to sociality is still unclear. Stress in the form of dominant audience members inhibits understanding of intentions, downgrading cognition to understanding of behaviour, but intentional communication may enable social bonding in stressful conditions. We examined the associations between gestural communication, sociality, stress and the outcome of interactions in wild chimpanzees. Social network size was positively associated with intentional but not non-intentional communication. When a dominant bystander was present with whom the recipient was weakly bonded, and gesturing was non-intentional, recipients produced avoidance response toward signallers to whom they were weakly bonded, indicating understanding of behaviour. Signallers used intentional gestures more frequently to recipients who were stressed, and intentional gestures evoked approach behaviour by the recipients, indicating understanding of intentionality. These results suggest that the presence of dominant bystanders is stressful, inhibiting understanding of intentionality. However, intentional gestures facilitate social bonding by allowing understanding of intentions. The cognitive skills underpinning intentional gestures may therefore play a key role in enabling primates to meet the demands of sociality. This article is part of the theme issue 'Cognition, communication and social bonds in primates'. [ABSTRACT FROM AUTHOR]

Simple Summary: Chimpanzees moving into sanctuary settings from captive research settings have often lived in smaller groups at their previous facilities. However, chimpanzees in the wild live in large multi-male, multi-female groups that provide them with a greater variety of social opportunities. Chimp Haven introduces chimpanzees to each other to form larger, more complex groups. This has resulted in hundreds of introductions or social integrations over a ten-year period. There are very limited published data on social integrations in chimpanzees. The purpose of the current study is to determine how group size, location, rearing, age, and sex differences affect the chimpanzees' affiliative, aggressive, and submissive behaviors during these social integrations to help inform future efforts at socializing chimpanzees in captive settings. The findings of the study are likely to assist us in determining which factors might be most important to consider for chimpanzee introductions. Chimp Haven is a sanctuary for chimpanzees being retired from biomedical research and from facilities that can no longer care for them. Chimpanzees often live in smaller groups in captive settings; however, Chimp Haven integrates them into larger, more species-typical groups. Social integrations, the process of introducing unfamiliar chimpanzees to one another, are often complex in terms of logistics and can be stressful due to the territorial nature of the animals, reduced space in captivity, and the fact that these situations are engineered by humans. From 2005 to 2015, Chimp Haven conducted 225 social integrations including 282 chimpanzees (male: n = 135; female: n = 147). Each integration involved 2 to 26 chimpanzees (mean = 9) and their age ranged from < one year old to 59 years old (mean = 30). We collected data ad libitum during the first 60 min after doors were opened between unfamiliar chimpanzees. The chimpanzees' affiliative, aggressive, and submissive behaviors were examined, comparing the subject's sex, rearing history, location/enclosure type, and group size impacts on these behaviors. The subject's sex, location, and group size were associated with the frequency of affiliative behaviors observed during social integration. All variables except for group size were associated with the frequency of aggressive behavior. The frequency of submissive behavior differed based on the subject's sex, rearing history, and group size. We were unable to make comparisons between successful and unsuccessful integrations, as most of these integrations were successful. [ABSTRACT FROM AUTHOR]

Simple Summary: A 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. [ABSTRACT FROM AUTHOR]

Objectives: In the human brain, the inferior frontal gyrus (IFG) is comprised of three morphological regions, which include the pars opercularis, pars triangularis, and pars orbitalis. These brains regions are implicated in a number of cognitive and linguistic functions, and the pars opercularis and pars triangularis of the language dominant hemisphere are collectively referred to as Broca's area. Evolution of the morphology of the IFG has been a topic of study in comparative neuroscience. Materials and Methods: Using magnetic resonance images (MRI) from 294 chimpanzees with known pedigrees, we quantified the folding patterns, surface area, and depth of three sulci and their subdivisions that define the IFG. Results: Chimpanzees show considerable individual variation in IFG sulci. A higher proportion of males and chimpanzees with better orofacial motor control had a bifurcation in the dorsal limb of the fronto‐orbital sulcus. For surface area and depth, chimpanzees with better orofacial motor control had greater leftward asymmetries in surface area and increased gyrification for the fronto‐orbital sulcus and greater rightward biases for the inferior precentral sulcus. Quantitative genetic analyses revealed that the average surface area and depth for three of the five sulci subdivisions were significantly heritable. By contrast, we found no evidence for heritability in asymmetries for any of the sulci. Discussion: These findings provide context to understanding evolutionary selection for increasing motor, cognitive and linguistic functions among primates, and contribute an essential comparative framework for interpreting IFG evolution in the human lineage. [ABSTRACT FROM AUTHOR]

Visitor effect studies have had inconsistent results, due in part to the inability to control for all confounding variables such as time of day, seasonal weather patterns, and so forth. This study represents the first instance where chimpanzee (Pan troglodytes) wounding frequencies were investigated across extensive time periods in the presence and complete absence of visitors, thus eliminating many visitor-related variables. Additional variables were eliminated through the zoo selection process, based on institutional responses to a 29-question survey, providing a novel approach to the question of visitor effects. The aim of this study was to determine if visitors were associated with a change in chimpanzee wound event frequencies across four 51-day time conditions, three of which occurred prior to the COVID-19 pandemic, and one during the first wave of pandemic-related zoo closures. We analyzed the archival records of 21 chimpanzees housed at three U.S. zoos. Due to the small number of wound events across all study windows, frequencies of "no wound" events were analyzed. A chi-square goodness of fit test was performed to determine whether the frequency of "no wound" events was equal between the "open" and "closed" trials. Results indicate that the frequencies did not differ, suggesting that chimpanzee welfare, as it relates to wounding, may not be adversely affected by zoo visitors. [ABSTRACT FROM AUTHOR]

Simple Summary: Environmental 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. [ABSTRACT FROM AUTHOR]

Simple Summary: To improve the lives of chimpanzees living in managed care, it is important to understand why their lives have ended. Previous efforts to document the causes of death for chimpanzees living in research facilities have been useful; however, interpretations need to consider the relatively small number of facilities assessed, the recency of those evaluations, and the possibility that mortality rates were affected by experimental interventions at these sites. Here, the causes of death of 224 chimpanzees at 42 accredited zoos over a 25-year period were analyzed. Most of the chimpanzees over 1 month of age died from causes associated with degenerative diseases (50%), infection (26%), and trauma (15%). Causes of fatal trauma ranged from aggression from conspecifics, the outcomes of exhibit escapes, or accidental drowning. The number of accidental deaths has decreased in the past decade, likely as a result of advances in exhibit design and management. Cardiopulmonary problems have long been known to be prevalent within the chimpanzee population, and they remain a major contributor to death for both sexes in this population. Understanding these findings provides an opportunity to improve the care and management of captive chimpanzees. Understanding causes of death allows adjustment of health management strategies for animals in managed care. From 224 documented chimpanzee deaths occurring from 1995 to 2019 in 42 accredited U.S. zoos, post-mortem records and necropsy reports were analyzed for the primary cause of death, which were available for 214 individuals. In total, 37 cases of stillbirth and neonatal deaths were assessed (16.5%); however, the focus was otherwise placed on the remaining 177 cases in which the death occurred in individuals aged greater than 1 month. There were no sex-related differences in etiology; however, age variation in the cause of death was statistically significant (p < 0.001). Elderly (35 years and older) chimpanzees tended to die of intrinsic, often degenerative, etiologies, whereas infants, juveniles, and adolescents (less than 15 years) were more likely to be involved in fatal trauma. Overall, there were 27 deaths (15.3% of all post-neonatal deaths) related to trauma and 13 of these were directly or indirectly related to conspecific aggression. Understanding causes of mortality and the interrelation with management can benefit managed populations of chimpanzees. [ABSTRACT FROM AUTHOR]

A large body of literature demonstrates the adaptive benefits of social relationships between kin, including fitness and survival. Given that most social mammals are characterized by male-biased dispersal, the majority of research on kin selection and associated advantages focuses on social relationships between female kin. Meanwhile, research on social relationships between adult male and female kin has primarily focused on inbreeding avoidance or the benefit to adult sons, with less attention on potential advantages these social relationships may provide females. The general pattern of male dominance over females in most mammal species suggests that females may benefit from protective associations with adult male kin. Using 43 years of behavioural data on the wild chimpanzees, Pan troglodytes schweinfurthii , of Gombe National Park, Tanzania, we examined association patterns between females and their adult maternal male kin. We specifically focused on how these associations may represent a trade-off between inbreeding avoidance and protection for females, particularly against infanticide. In accordance with inbreeding avoidance, we predicted that females' association with adult kin would decrease when they were maximally tumescent, signalling sexual receptivity. To determine whether female–male kin associations provide protection to females, we examined female associations with adult male kin during their first year postpartum when infants are most vulnerable to infanticide. We predicted that during this first year postpartum, females would have a higher association with male kin than with unrelated males. We found that females associated more with adult sons and brothers than with unrelated males when they did not have a sexual swelling. Female association increased with all males across tumescence but females associated less with their brothers than they did with their sons and unrelated males when they were maximally tumescent, providing equivocal support for the inbreeding avoidance hypothesis. Furthermore, females associated more with both sons and brothers than with unrelated males in the first 6 months of the postpartum period. Higher association with brothers, relative to unrelated males, persisted throughout the first year postpartum. Together, these results speak to the cost–benefit trade-off in female and adult male kin associations, highlighting the potential protective advantage for females, especially during the postpartum period. • We examined association patterns between females and their adult maternal male kin. • Association with sons, brothers and unrelated males increased with swelling size. • Association with sons was higher than with other males during early postpartum. • Association with brothers was higher than with unrelated males 1 year postpartum. • Our results suggest a potential male kin protective advantage postpartum. [ABSTRACT FROM AUTHOR]

An influential view in primate social cognition research is that chimpanzees, as well as other great apes, attribute the mental relation of seeing to others. In a recent article, Tyler Burge challenges this view and argues that data from studies with chimpanzees provide no evidence that chimpanzees attribute the mental relation of seeing but only that they attribute the nonmental relation of sensing. Burge's argument rests upon the claim that attributions of seeing attribute more than attributions of sensing. We contend that the reasons Burge offers for this claim are unconvincing. We go on to argue that data from a recent study with chimpanzees cannot be explained on the hypothesis that chimpanzees attribute sensing but can be explained on the hypothesis that they attribute seeing. We conclude that, as things stand, there is currently more reason to take chimpanzees to attribute the mental relation of seeing than the nonmental relation of sensing. [ABSTRACT FROM AUTHOR]

In the deltoid muscles of Pan troglodytes and Homo sapiens, we have analyzed the muscle architecture and the expression of the myosin heavy chain (MHC) isoforms. Our aim was to identify differences between the two species that could be related to their different uses of the upper limb. The deltoid muscle of six adult Pan troglodytes and six adult Homo sapiens were dissected. The muscle fascicle length (MFL) and the physiological cross‐sectional area (PCSA) of each muscle were calculated in absolute and normalized values. The expression pattern of the MHC‐I, MHC‐IIa and MHC‐IIx isoforms was analyzed in the same muscles by real‐time polymerase chain reaction. Only the acromial deltoid (AD) presented significant architectural differences between the two species, with higher MFL values in humans and higher PCSA values in chimpanzees. No significant differences in the expression pattern of the MHC isoforms were identified. The higher PCSA values in the AD of Pan troglodytes indicate a greater capacity of force generation in chimpanzees than in humans, which may be related to a greater use of the upper limb in locomotion, specifically in arboreal locomotion like vertical climbing. The functional differences between chimpanzees and humans in the deltoid muscle are more related to muscle architecture than to a differential expression of MHC isoforms. [ABSTRACT FROM AUTHOR]

A fundamental aspect of human communication is our ability to refer to external objects and events through both words and gestures (such as pointing), yet the evolutionary origins of such signals remain obscure. Apes, living in their natural environments, rarely or never point, but it has been claimed that male chimpanzees, Pan troglodytes schweinfurthii , from the Ngogo community, Uganda, habitually use exaggerated loud scratches (ELSs) to refer to specific body locations where they wish to be groomed (Pika & Mitani, 2006, Current Biology , 16 (6), 191–192). This study suggested continuity between referential abilities in humans and our closest living relatives, making it an important finding to replicate in other populations. Hence here, we compared whether ELSs are used in a referential manner across four wild communities of eastern chimpanzees (Ngogo, Kanyawara, Sonso and Waibira). Our data show that scratchers were significantly more likely to receive grooming in the scratched location at Ngogo compared to the other three sites. At the latter sites this response occurred at low rates and signallers did not seem to pursue this goal. This suggests that ELSs do not function referentially at these sites, and the published findings from Ngogo were not replicated. Further exploration into alternative functions of ELSs in the Kanyawara community revealed that, in this community, this signal functions to initiate grooming bouts and to reengage partners during grooming pauses. Individuals who produced the signal to initiate grooming were likely to offer grooming. In contrast, during grooming bouts, groomers produced ELSs to request reciprocation of grooming from their partner. Our study demonstrates that chimpanzees do not ubiquitously use the ELS in a referential manner, but that they can use this gesture in a highly flexible fashion, with signal function depending on the intricate details of the social contexts in which they are produced. • Previous findings of referential gesture use in wild chimpanzees are not replicated. • Habitual use of referential gestures is not ubiquitous in wild chimpanzees. • In one community exaggerated loud scratches (ELSs) fulfil two alternative functions. • ELSs are used to initiate grooming bouts and to request grooming reciprocation. • ELSs play a role in negotiating the dynamics of cooperative grooming interactions. [ABSTRACT FROM AUTHOR]

Human between-group interactions are highly variable, ranging from violent to tolerant and affiliative. Tolerance between groups is linked to our unique capacity for large-scale cooperation and cumulative culture, but its evolutionary origins are understudied. In chimpanzees, one of our closest living relatives, predominantly hostile between-group interactions impede cooperation and information flow across groups. In contrast, in our other closest living relative, the bonobo, tolerant between-group associations are observed. However, as these associations can be frequent and prolonged and involve social interactions that mirror those within groups, it is unclear whether these bonobos really do belong to separate groups. Alternatively, the bonobo grouping patterns may be homologous to observations from the large Ngogo chimpanzee community, where individuals form within-group neighborhoods despite sharing the same membership in the larger group. To characterize bonobo grouping patterns, we compare the social structure of the Kokolopori bonobos with the chimpanzee group of Ngogo. Using cluster analysis, we find temporally stable clusters only in bonobos. Despite the large spatial overlap and frequent interactions between the bonobo clusters, we identified significant association preference within but not between clusters and a unique space use of each cluster. Although bonobo associations are flexible (i.e., fission-fusion dynamics), cluster membership predicted the bonobo fission compositions and the spatial cohesion of individuals during encounters. These findings suggest the presence of a social system that combines clear in-group/out-group distinction and out-group tolerance in bonobos, offering a unique referential model for the evolution of tolerant between-group interactions in humans. [ABSTRACT FROM AUTHOR]

Interpretations of values of the FST measure of genetic differentiation rely on an understanding of its mathematical constraints. Previously, it has been shown that FST values computed from a biallelic locus in a set of multiple populations and FST values computed from a multiallelic locus in a pair of populations are mathematically constrained as a function of the frequency of the allele that is most frequent across populations. We generalize from these cases to report here the mathematical constraint on FST given the frequency M of the most frequent allele at a multiallelic locus in a set of multiple populations. Using coalescent simulations of an island model of migration with an infinitely-many-alleles mutation model, we argue that the joint distribution of FST and M helps in disentangling the separate influences of mutation and migration on FST. Finally, we show that our results explain a puzzling pattern of microsatellite differentiation: the lower FST in an interspecific comparison between humans and chimpanzees than in the comparison of chimpanzee populations. We discuss the implications of our results for the use of FST. [ABSTRACT FROM AUTHOR]

Areas dedicated to higher brain functions such as the orbitofrontal cortex (OFC) are thought to be unique to hominidae. The OFC is involved in social behavior, reward and punishment encoding and emotional control. Here, we focused on the putative corresponding area in the sheep to assess its homology to the OFC in humans. We used classical histology in five sheep (Ovis aries) and four chimpanzees (Pan troglodytes) as a six-layered-cortex primate, and Diffusion Tensor Imaging (DTI) in three sheep and five human brains. Nissl's staining exhibited a certain alteration in cortical lamination since no layer IV was found in the sheep. A reduction of the total cortical thickness was also evident together with a reduction of the prevalence of layer one and an increased layer two on the total thickness. Tractography of the sheep OFC, on the other hand, revealed similarities both with human tracts and those described in the literature, as well as a higher number of cortico-cortical fibers connecting the OFC with the visual areas in the right hemisphere. Our results evidenced the presence of the basic components necessary for complex abstract thought in the sheep and a pronounced laterality, often associated with greater efficiency of a certain function, suggested an evolutionary adaptation of this prey species. [ABSTRACT FROM AUTHOR]

Objectives: In the present study, we have analyzed the anatomy of the radiocarpal joint ligaments and muscles in Pan troglodytes and Homo sapiens in order to identify similarities and differences between the two species that may be related to differences in hand use and function. Materials and Methods: Anatomical dissections of the ligaments and muscles of the radiocarpal joint were performed in six adult chimpanzees and 12 humans. The mass of each ligament and of the functional ligament groups were calculated relative to the total ligament mass and compared between the two species. The mass of the functional muscle groups relative to the total mass of the muscles of the radiocarpal joint was also calculated and compared between the two species. Results: The ligaments of the radiocarpal joint had similar anatomical characteristics in chimpanzees and humans. The relative mass of the palmar ligaments was greater in humans, while that of the dorsal radiocarpal ligament was greater in chimpanzees. In both species, the relative mass of the palmar and dorsal muscle groups was inversely related to that of the corresponding ligament groups. Discussion: The greater relative mass of the palmar ligaments in humans may be related to the importance of wrist extension during manipulative tasks. The greater relative mass of the dorsal radiocarpal ligament in chimpanzees may be related to the need to stabilize the radiocarpal joint in flexion, mainly during arboreal locomotion. [ABSTRACT FROM AUTHOR]

Previous studies have revealed that non-human primates can differentiate the age category of faces. However, the knowledge about age recognition in non-human primates is very limited and whether non-human primates can process facial age information in a similar way to humans is unknown. As humans have an association between time and space (e.g., a person in an earlier life stage to the left and a person in a later life stage to the right), we investigated whether chimpanzees spatially represent conspecifics' adult and infant faces. Chimpanzees were tested using an identical matching-to-sample task with conspecific adult and infant face stimuli. Two comparison images were presented vertically (Experiment 1) or horizontally (Experiment 2). We analyzed whether the response time was influenced by the position and age category of the target stimuli, but there was no evidence of correspondence between space and adult/infant faces. Thus, evidence of the spatial representation of the age category was not found. However, we did find that the response time was consistently faster when they discriminated between adult faces than when they discriminated between infant faces in both experiments. This result is in line with a series of human face studies that suggest the existence of an "own-age bias." As far as we know, this is the first report of asymmetric face processing efficiency between infant and adult faces in non-human primates. [ABSTRACT FROM AUTHOR]

It is known that methods to estimate the rate of adaptive evolution, which are based on the McDonald–Kreitman test, can be biased by changes in effective population size. Here, we demonstrate theoretically that changes in population size can also generate an artifactual correlation between the rate of adaptive evolution and any factor that is correlated to the strength of selection acting against deleterious mutations. In this context, we have investigated whether several site-level factors influence the rate of adaptive evolution in the divergence of humans and chimpanzees, two species that have been inferred to have undergone population size contraction since they diverged. We find that the rate of adaptive evolution, relative to the rate of mutation, is higher for more exposed amino acids, lower for amino acid pairs that are more dissimilar in terms of their polarity, volume, and lower for amino acid pairs that are subject to stronger purifying selection, as measured by the ratio of the numbers of nonsynonymous to synonymous polymorphisms (p N/ p S). All of these correlations are opposite to the artifactual correlations expected under contracting population size. We therefore conclude that these correlations are genuine. [ABSTRACT FROM AUTHOR]

Viral infection is a major cause of ill health in wild chimpanzees (Pan troglodytes), but most evidence to date has come from conspicuous disease outbreaks with high morbidity and mortality. To examine the relationship between viral infection and ill health during periods not associated with disease outbreaks, we conducted a longitudinal study of wild eastern chimpanzees (P. t. schweinfurthii) in the Kanyawara and Ngogo communities of Kibale National Park, Uganda. We collected standardized, observational health data for 4 years and then used metagenomics to characterize gastrointestinal viromes (i.e., all viruses recovered from fecal samples) in individual chimpanzees before and during episodes of clinical disease. We restricted our analyses to viruses thought to infect mammals or primarily associated with mammals, discarding viruses associated with nonmammalian hosts. We found 18 viruses (nine of which were previously identified in this population) from at least five viral families. Viral richness (number of viruses per sample) did not vary by health status. By contrast, total viral load (normalized proportion of sequences mapping to viruses) was significantly higher in ill individuals compared with healthy individuals. Furthermore, when ill, Kanyawara chimpanzees exhibited higher viral loads than Ngogo chimpanzees, and males, but not females, exhibited higher infection rates with certain viruses and higher total viral loads as they aged. Post‐hoc analyses, including the use of a machine‐learning classification method, indicated that one virus, salivirus (Picornaviridae), was the main contributor to health‐related and community‐level variation in viral loads. Another virus, chimpanzee stool‐associated virus (chisavirus; unclassified Picornavirales), was associated with ill health at Ngogo but not at Kanyawara. Chisavirus, chimpanzee adenovirus (Adenoviridae), and bufavirus (Parvoviridae) were also associated with increased age in males. Associations with sex and age are consistent with the hypothesis that nonlethal viral infections cumulatively reflect or contribute to senescence in long‐lived species such as chimpanzees. Research Highlights: Some viruses of wild primates are relatively benign but may be useful as biomarkers of physiology and health.We used metagenomic methods to characterize viral communities in feces of wild chimpanzees in two long‐term study communities (Kanyawara and Ngogo) in Kibale National Park, Uganda.Chimpanzees exhibited higher viral loads (normalized abundances of viral genetic sequences) when they were ill than when those same individuals were previously healthy.When ill, Kanyawara chimpanzees exhibited higher viral loads than Ngogo chimpanzees, and males, but not females, exhibited higher viral loads as they aged. [ABSTRACT FROM AUTHOR]