Chimpanzees Research Articles

Compassion Fatigue in Chimpanzee (Pan troglodytes) Caregivers: Prevalence, Contributing Factors, and Coping Mechanisms

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.

Testing the reliability of the rearticulation of osteological primate pelves in comparative morphological studies.

The evolution of human pelvic form is primarily studied using disarticulated osteological material of living and fossil primates that need rearticulation to approximate anatomical position. To test whether this technique introduces errors that impact biological signals, virtual rearticulations of the pelvis in anatomical position from computed tomography scans were compared with rearticulated models from the same individuals for one female and one male of Homo sapiens, Pan troglodytes, Macaca mulatta, Lepilemur mustelinus, Galago senegalensis, and Nycticebus pygmaeus. "Cadaveric" pelvic bones were first analyzed in anatomical position, then the three bones were segmented individually, intentionally scattered, and "rearticulated" to test for rearticulation error. Three-dimensional landmarks and linear measurements were used to characterize the overall pelvis shape. Cadaveric and rearticulated pelves were not identical, but inter-specific and intra-specific shape differences were higher than the landmarking error in the cadaveric individuals and the landmarking/rearticulation error in the rearticulated pelves, demonstrating that the biological signal is stronger than the noise introduced by landmarking and rearticulation. The rearticulation process, however, underestimates the medio-lateral pelvic measurements in species with a substantial pubic gap (e.g., G. senegalensis, N. pygmaeus) possibly because the greater contribution of soft tissue to the pelvic girdle introduces higher uncertainty during rearticulation. Nevertheless, this discrepancy affects only the caudal-most part of the pelvis. This study demonstrates that the rearticulation of pelvic bones does not substantially affect the biological signal in comparative 3D morphological studies but suggests that anatomically connected pelves of species with wide pubic gaps should be preferentially included in these studies.

Sketches of chimpanzee (Pan troglodytes) hoo’s: vowels by any other name?

In human speech, the close back rounded vowel /u/ (the vowel in “boot”) is articulated with the tongue arched toward the dorsal boundary of the hard palate, with the pharyngeal cavity open. Acoustic and perceptual properties of chimpanzee (Pan troglodytes) hoo’s are similar to those of the human vowel /u/. However, the vocal tract morphology of chimpanzees likely limits their phonetic capabilities, so that it is unlikely, or even impossible, that their articulation is comparable to that of a human. To determine how qualities of the vowel /u/ may be achieved given the chimpanzee vocal tract, we calculated transfer functions of the vocal tract area for tube models of vocal tract configurations in which vocal tract length, length and area of a laryngeal air sac simulacrum, length of lip protrusion, and area of lip opening were systematically varied. The method described is principally acoustic; we make no claim as to the actual shape of the chimpanzee vocal tract during call production. Nonetheless, we demonstrate that it may be possible to achieve the acoustic and perceptual qualities of back vowels without a reconfigured human vocal tract. The results, while tentative, suggest that the production of hoo’s by chimpanzees, while achieving comparable vowel-like qualities to the human /u/, may involve articulatory gestures that are beyond the range of the human articulators. The purpose of this study was to (1) stimulate further simulation research on great ape articulation, and (2) show that apparently vowel-like phenomena in nature are not necessarily indicative of evolutionary continuity per se.

First detection of Rift Valley fever virus antibodies in non-human primates in Cameroon.

We tested for Rift Valley fever virus (RVFV) from at least 15 species of non-human primates. RVFV IgG/IgM antibodies were detected in 3.7% (2 out of 53) of chimpanzees (Pan troglodytes) and in 1.4% (1 out of 72) of unidentified non-human primate species. This study was the first investigation of RVFV in monkeys in Cameroon.

Dental findings in wild great apes from macerated skull analysis.

Oral health is a crucial aspect of overall well-being in both humans and nonhuman primates. Understanding the oral pathologies and dental conditions in apes can provide valuable insights into their evolutionary history, dietary habits, and overall health. The present study evaluates dental findings in wild great apes from museum specimens to gain insights into the influence of natural nutrition on dental health. Complete macerated skulls of wild, adult great apes from the collection of the Museum of Natural History, Berlin, Germany, were examined. We analyzed skulls of 53 gorillas (Gorilla gorilla), 63 chimpanzees (Pan troglodytes), and 41 orangutans (Pongo spp.). For each skull, we recorded wear of dental hard tissues (Lussi and Ganss index), carious lesions, and periodontal bone loss. Incisal and occlusal dental hard tissue defects were found in all skulls, as well as considerable external staining. In all species, incisors and canines showed the greatest loss of tissue, followed by molars. The wear of molars decreased from the first to the third molars, premolars showed the least pronounced defects. Some individuals had apical osteolytic defects along with severe dental hard tissue loss with pulp involvement or after dental trauma, respectively (n = 5). Our study did not observe any carious lesions among the examined great ape skulls. However, we did find evidence for localized or generalized periodontal bone loss in a subset of the specimens (n = 3 chimpanzees, n = 7 orangutans). The natural diet and foraging behavior of great apes induces abrasion and attrition of dental hard tissue but does not yield carious lesions. The occurrence of periodontitis in individual apes indicates that the natural circumstances can induce periodontal bone loss even in the wild, despite physiological nutrition.

Instrumental helping and short‐term reciprocity in chimpanzees and human children

AbstractChimpanzees (Pan troglodytes) and humans cooperate in reciprocal patterns, but it is unclear whether these interactions are based on the same psychological foundations. While there is evidence suggesting that both species engage in long‐term forms of reciprocity, there is very little work exploring their short‐term behavioural contingencies with suitable methods. Here, we present a direct comparative study on short‐term reciprocity in chimpanzees and human children using a novel, low‐cost instrumental helping task. We investigated whether participants help a conspecific partner to obtain a tool for accessing a reward, and whether the level of helping depends on the partner's previous helpful or unhelpful behaviour. In line with prior research, both chimpanzees and children demonstrated helping behaviour towards their partner. However, the extent to which the two species showed short‐term reciprocity differed considerably. After receiving help, tested children always helped in return. They helped substantially less when interacting with an unhelpful partner. Chimpanzees showed a higher tendency to help when interacting with a helpful compared to an unhelpful partner only in the first half the experiment. With increasing trial number, chimpanzees stopped discriminating between helpful and unhelpful partners. This study provides evidence for short‐term reciprocity in human children and, to a lesser extent, in our closest living relatives. Our findings demonstrate that helping paradigms provide a useful context to investigate reciprocal motives in humans and chimpanzees alike.

Cross-species amplification from non-human primate DNAs in commercial human DNA assay kits

Species specificity of commercial human DNA quantification kits and short tandem repeat (STR) profiling kits was examined using primate DNA samples. These samples comprised 33 individuals from eight primate species, each with gender and kinship data, including human (Homo sapiens), chimpanzee (Pan troglodytes), gorilla (Gorilla gorilla), and orangutan (Pongo pygmaeus) of Hominidae family, and Japanese macaque (Macaca fuscata), long-tailed macaque (Macaca fascicularis), hamadryas baboon (Papio hamadryas), and savannah monkey (Chlorocebus sp.) of Cercopithecidae family. The findings revealed varying levels of cross-species amplifications in all non-human DNA samples that correlated with their evolutionary proximity to humans, both kit types. Moreover, cross-species amplification, including female DNA samples, was observed in a Y-chromosomal STR profiling kit. Additionally, species specificity differed among the commercial kits examined. The cross-species amplification data presented in this study offer valuable assistance in interpreting the results of individual human identification in forensic cases involving non-human primates.

Evaluating modularity in the hominine skull related to feeding biomechanics.

Modular architecture of traits in complex organisms can be important for morphological evolution at micro- and sometimes macroevolutionary scales as it may influence the tempo and direction of changes to groups of traits that are essential for particular functions, including food acquisition and processing. We tested several distinct hypotheses about craniofacial modularity in the hominine skull in relation to feeding biomechanics. First, we formulated hypothesized functional modules for craniofacial traits reflecting specific demands of feeding biomechanics (e.g., masseter leverage/gape or tooth crown mechanics) in Homo sapiens, Pan troglodytes, and Gorilla gorilla. Then, the pattern and strength of modular signal was quantified by the covariance ratio coefficient and compared across groups using covariance ratio effect size. Hierarchical clustering analysis was then conducted to examine whether a priori-defined functional modules correspond to empirically recovered clusters. There was statistical support for most a priori-defined functional modules in the cranium and half of the functional modules in the mandible. Modularity signal was similar in the cranium and mandible, and across the three taxa. Despite a similar strength of modularity, the empirically recovered clusters do not map perfectly onto our priori functional modules, indicating that further work is needed to refine our hypothesized functional modules. The results suggest that modular structure of traits in association with feeding biomechanics were mostly shared with humans and the two African apes. Thus, conserved patterns of functional modularity may have facilitated evolutionary changes to the skull during human evolution.

Facial Expression of TIPI Personality and CHMP-Tri Psychopathy Traits in Chimpanzees (Pan troglodytes)

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.

High urinary oxidative DNA damage in wild chimpanzees ranging in proximity of agricultural fields in Sebitoli area, Uganda.

Agriculture expansion is a major cause of habitat loss and exposure to phytochemical pollution for non-human primates. In addition to endocrine disruption, exposure to pesticides may have other sublethal physiological consequences for animals, such as generation of oxidative damage to macromolecules. In this study, we analyzed the pesticides contained in the river water across the home range of wild chimpanzees (Pan troglodytes) in Sebitoli area located on the Northern part of Kibale National Park (Uganda). We tested whether levels of three urinary markers of oxidative damage vary among individuals in relation to their ranging patterns, as a proxy for pesticide exposure intensity. To better characterize the foraging habitat use, the trophic level, and the energetic status of study individuals, we also quantified urinary levels of carbon and nitrogen stable isotope signatures and of C-peptide. Among the 511 pesticides screened, 18 compounds including herbicides, insecticides, and fungicides were found in the water sampled in the Western part of the home range of chimpanzees. In this area, chimpanzees used to feed on maize crops. By contrast, in the Eastern part where crop feeding was never observed, we found only seven pesticides. According to their ranging patterns and thus crop feeding frequency, the 139 urine samples collected from 43 Sebitoli chimpanzees were categorized as belonging to low, medium, and high exposure level. Chimpanzees from the high exposure zone had higher oxidative DNA damage (8-OHdG) than chimpanzees from both the low and medium exposure groups, who had similar levels of oxidative DNA damage. In addition, individuals with higher C-peptide tended to have significantly higher oxidative DNA damage and lipid peroxides. The three exposure groups had similar levels of urinary 8-isoprostanes and of urinary lipid peroxides. These results were robust for any potential confounding effect of other variables because neither age category nor sex or isotope levels were significantly associated with markers of oxidative damage. Our study points to genotoxic effects as one potential sublethal consequence of ranging in proximity of agricultural fields owing to exposure to pesticides or other unidentified sources of stress. Given our phylogenetic proximity, this information is relevant for the conservation of this species which is endangered and also sentinel for human health.

Investigating infant feeding development in wild chimpanzees using stable isotopes of naturally shed hair.

Measuring the relative contributions of milk and non-milk foods in the diets of primate infants is difficult from observations. Stable carbon (δ13 C) and nitrogen (δ15 N) isotopes in hair can be used to physiologically track infant feeding through development, but few wild studies have done so, likely due to the difficulty in collecting hair non-invasively. We assessed infant feeding at different ages in wild chimpanzees (Pan troglodytes) at Ngogo, Uganda using δ13 C and δ15 N of keratin in 164 naturally shed hairs from 29 infants (61 hairs), 6 juveniles (7 hairs), 28 mothers (67 hairs) and 14 adult males (29 hairs). Hairs were collected when they stuck to feces during defecation or from the ground after chimpanzees groomed or rested. We could not distinguish between the hairs of infants and mothers using strand length and diameter. Infants 1-2 years old were most enriched in 13 C and 15 N and showed means of 1.1‰ in δ13 C and 2.1‰ in δ15 N above their mothers. Infants at 2 years had hair δ13 C values like those of their mothers, which suggests infants began relying more heavily on plants around this age. While mother-infant δ13 C and δ15 N differences generally decreased with offspring age, as is expected when infants rely increasingly more on independent foraging through development, milk seemed to remain an important dietary component for infants older than 2.5 years, as evidenced by continuing elevated δ15 N. We showed that stable carbon and nitrogen isotopes in naturally shed hairs can feasibly detect trophic level differences between chimpanzee infants and mothers. Since it can mitigate some of the limitations associated with behavioral and fecal stable isotope data, the use of hair stable isotopes is a useful, non-invasive tool for assessing infant feeding development in wild primates.

Trends and biases in research efforts for primate conservation: threatened species are not in the spotlight

The order Primates is a diverse group with worrisome conservation status, in which 67% of the species are threatened, and 85% have declining populations. Although the studies in primate conservation have increased over the past two decades, there is a lack of knowledge about the trends and biases in these conservation research efforts. We reviewed the primate conservation literature to identify the trends in allocating research efforts across species, themes, and countries. We also investigated whether the studies are biased by primate richness, species body mass, range size, locomotion type, diel activity, threat status, time since species description, and phylogenetic relatedness. We found that the highest number of studies was about habitat fragmentation. Madagascar, Indonesia, and Brazil concentrated most of the studies. Pan troglodytes was the most studied species. The conservation research efforts are skewed towards primate-rich countries, earlier-described and large species that use arboreal and terrestrial substrates, and that are phylogenetically related. Therefore, research in primate conservation seems more motivated by specific primate attributes rather than aspects of species vulnerability and their main threats. The elucidation of these trends and biases may help identify knowledge gaps and new research opportunities, contributing to optimizing future conservation research efforts in primate conservation.

Serum vitamin D in sanctuary chimpanzees (Pan troglodytes) in range countries: A pilot study.

Vitamin D is essential for skeletal health, calcium homeostasis and general health. The major and more stable form of vitamin D in circulation is 25-hydroxyvitamin D (25-OH-D); this is the most valuable indicator of vitamin D status. There are studies on laboratory and zoo-housed chimpanzees; however, serum vitamin D status has not been documented in chimpanzees in range countries. (1) Determine the range of circulating 25-OH-D concentrations in chimpanzees in range countries. (2) Assess the influence of age, sex, and sun exposure on 25-OH-D serum concentrations. Opportunistic blood samples were obtained from 127 clinically healthy chimpanzees. Serum 25-OH-D concentration was measured with a commercially available competitive ELISA. The median overall 25-OH-D concentration for chimpanzees in range countries was 46.24nmol/L (range: 17.10-109.23nmol/L). Males had a significantly lower concentration (40.15nmol/L) than females (49.61nmol/L), and infants (37.99nmol/L) had a significantly lower concentration than adults (46.04nmol/L). Concentrations of 25-OH-D in chimpanzees in sunnier habitats were significantly higher compared to thick tropical forest habitat. The present constitutes a large dataset of serum 25-OH-D concentrations in range country sanctuary chimpanzees and contributes to document normal ranges. Age, sex, and sun exposure influenced serum concentrations of 25-OH-D in sanctuary chimpanzees.

Covariation between the shape and mineralized tissues of the rib cross section in Homo sapiens, Pan troglodytes and Sts 14.

Studying rib torsion is crucial for understanding the evolution of the hominid ribcage. Interestingly, there are variables of the rib cross section that could be associated with rib torsion and, consequently, with the morphology of the thorax. The aim of this research is to conduct a comparative study of the shape and mineralized tissues of the rib cross section in different hominids to test for significant differences and, if possible, associate them to different thoracic morphotypes. The sample consists of the rib cross sections at the midshaft taken from 10 Homo sapiens and 10 Pan troglodytes adult individuals, as well as from A. africanus Sts 14. The shape of these rib cross sections was quantified using geometric morphometrics, while the mineralized tissues were evaluated using the compartmentalization index. Subsequently, covariation between both parameters was tested by a Spearman's ρ test, a permutation test and a linear regression. Generally, P. troglodytes individuals exhibit rib cross sections that are rounder and more mineralized compared to those of H. sapiens. However, the covariation between both parameters was only observed in typical ribs (levels 3-10). Although covariation was not found in the rib cross sections of Sts 14, their parameters are closer to P. troglodytes. On the one hand, the differences observed in the rib cross sections between H. sapiens and P. troglodytes might be related to different degrees of rib torsion and, consequently, to different thoracic 3D configurations. These findings can be functionally explained by considering their distinct modes of breathing and locomotion. On the other hand, although the rib cross sections belonging to Sts 14 are more similar to those of P. troglodytes, previous publications determined that their overall morphology is closer to modern humans. This discrepancy could reflect a diversity of post-cranial adaptations in Australopithecus.

Downclimbing and the evolution of ape forelimb morphologies.

The forelimbs of hominoid primates (apes) are decidedly more flexible than those of monkeys, especially at the shoulder, elbow and wrist joints. It is tempting to link the greater mobility of these joints to the functional demands of vertical climbing and below-branch suspension, but field-based kinematic studies have found few differences between chimpanzees and monkeys when comparing forelimb excursion angles during vertical ascent (upclimbing). There is, however, a strong theoretical argument for focusing instead on vertical descent (downclimbing), which motivated us to quantify the effects of climbing directionality on the forelimb kinematics of wild chimpanzees (Pan troglodytes) and sooty mangabeys (Cercocebus atys). We found that the shoulders and elbows of chimpanzees and sooty mangabeys subtended larger joint angles during bouts of downclimbing, and that the magnitude of this difference was greatest among chimpanzees. Our results cast new light on the functional importance of downclimbing, while also burnishing functional hypotheses that emphasize the role of vertical climbing during the evolution of apes, including the human lineage.

A non-invasive, concealed electrocardiogram and bioimpedance measurement system for captive primates.

Captive housed non-human primates, specifically great apes such as chimpanzees (Pan troglodytes) are frequently reported to have died from or are diagnosed with potentially fatal heart conditions that require the monitoring of physiological signals such as electrocardiogram (ECG) or respiratory rate. ECG screening must be conducted after applying full anaesthesia, causing potential physical and emotional stress as well as risk for the animal. Here, we present an electronic system that simultaneously measures the ECG and the electrical bioimpedance for the early detection of abnormal cardiovascular activity. Modified gloves whose fingers are equipped with electrodes enable the caregiver to obtain three cardiovascular signals (ECG, pulse rate and respiratory rate) by placing the fingertips on specific parts of the non-human primate without needing any prior physical preparations. Validation (ECG and bioimpedance) was performed both on humans and on captive housed chimpanzees, where all the signals of interest were correctly acquired.

Threat vocalisations are acoustically similar between humans (Homo sapiens) and chimpanzees (Pan troglodytes)

ABSTRACT In behavioural contexts like fighting, eating, and playing, acoustically distinctive vocalisations are produced across many mammalian species. Such expressions may be conserved in evolution, pointing to the possibility of acoustic regularities in the vocalisations of phylogenetically related species. Here, we test this hypothesis by comparing the degree of acoustic similarity between human and chimpanzee vocalisations produced in 10 similar behavioural contexts. We use two complementary analysis methods: Pairwise acoustic distance measures and acoustic separability metrics based on unsupervised learning algorithms. Cross-context analysis revealed that acoustic features of vocalisations produced when threatening another individual were distinct from other types of vocalisations and highly similar across species. Using a multimethod approach, these findings demonstrate that human vocalisations produced when threatening another person are acoustically similar to chimpanzee vocalisations in the same situation as compared to other types of vocalisations, likely reflecting a phylogenetically ancient vocal signalling system.

Functional morphological integration related to feeding biomechanics in the hominine skull

Quantifying and characterizing the pattern of trait covariances is crucial for understanding how population-level patterns of integration might constrain or facilitate craniofacial evolution related to the feeding system. This study addresses an important gap in our knowledge by investigating magnitudes and patterns of morphological integration of biomechanically informative traits in the skulls of Homo sapiens, Pan troglodytes, and Gorilla gorilla. We predicted a lower magnitude of integration among human biomechanical traits since humans eat a softer, less biomechanically challenging diet than apes. Indeed, compared to African apes, the magnitudes of integration were lower in H. sapiens skulls for form data (raw dimensions) but were similar or higher for shape data (raw dimensions scaled by geometric mean). Patterns of morphological integration were generally similar, but not identical, across the three species, particularly for the form data compared to the shape data. Traits that load heavily on the primary axis of variation in morphospace are generally associated with size and/or shape of the temporalis and masseter muscles and with dimensions related to the constrained lever model of jaw biomechanics. Given the conserved nature of morphological integration, skull adaptations for food processing in African apes and humans may have been constrained to occur along certain paths of high evolvability. The conserved pattern of functional integration also indicates that extant hominine species can operate as reasonable analogues for extinct hominins in studies that require population-level patterns of trait variance/covariance.

A case of intercommunity lethal aggression by chimpanzees in an open and dry landscape, Issa Valley, western Tanzania

Intercommunity (lethal) aggression is a familiar component of the behavioural repertoire of many forest-dwelling chimpanzee (Pan troglodytes) communities. However, until now, the absence of intercommunity attacks – including killings – in communities that live in open, mosaic environments has supported hypotheses of reduced resource competition in drier habitats, and informed referential models of early hominin social dynamics in a similar habitat. In June 2020, we observed the first instance of intercommunity lethal aggression, a male-committed infanticide, by the Issa chimpanzee community, which live in a savannah-mosaic habitat in the Issa Valley, western Tanzania. The carcass was recovered by researchers after it was abandoned by the attackers. Here, we give a detailed account of the events leading up to and including the infanticide, and contextualise our observations with what has been described for other chimpanzee communities. Notably, in contrast to the majority of reported intercommunity infanticides, the infant male victim was castrated (and not cannibalised), making this the youngest reported castration. This observation of intercommunity aggression disproves its hypothesised absence in savannah-dwelling chimpanzees, which by extension, has implications for early hominin evolution. We suggest that the near absence of observations of intercommunity aggression in savannah chimpanzee communities is most likely due to the lack of long-term study communities, and in some cases geographic isolation. We hypothesise that food-rich areas within a habitat with otherwise widely distributed food sources may select for intense intercommunity aggression despite the low population density characteristic of savannah communities. Anecdotes such as this add to the comparative database available on intercommunity killings in chimpanzee society, improving our ability to draw inferences about their evolutionary significance.

Behavioral Welfare Research for the Management of Sanctuary Chimpanzees (Pan troglodytes).

Chimp Haven is a sanctuary for chimpanzees retired from biomedical research, rescued from the pet trade, or re-homed after other organizations could no longer care for them. To provide optimal care for over 300 chimpanzees, Chimp Haven's animal care team includes experts in behavioral science, veterinary treatment, and husbandry practices. To aid these teams in making routine welfare management decisions, a system of behavioral metrics provides objective data to guide decisions and track outcomes. Chimp Haven has built and piloted seven behavioral metric protocols over the past 5 years to provide staff with an objective and comprehensive picture of the chimpanzees' behavioral welfare. The data from behavioral observations, staff surveys, and routine staff documentation are analyzed and processed through Google Forms, ZooMonitor, Microsoft Power Bi, Microsoft Excel, and R. Each metric assists staff in making data-based decisions regarding the management of captive chimpanzees related to abnormal behavior, hair loss, wounding, social relationships, positive reinforcement training and overall wellness. In this article, we explore examples of each metric and how they have been utilized to monitor and make decisions for both social groups of chimpanzees as well as individuals. These metrics can be collected and shared easily in an understandable format, which may provide an important framework for others to follow to enable the tracking of welfare for other sanctuaries, non-human primates, as well as other species.