Fiber-type phenotype of the jaw-closing muscles in Gorilla gorilla, Pan troglodytes, and Pan paniscus: A test of the Frequent Recruitment Hypothesis.

Skeletal muscle fiber types are important determinants of the contractile properties of muscle fibers, such as fatigue resistance and shortening velocity. Yet little is known about how jaw-adductor fiber types correlate with feeding behavior in primates. Compared with chimpanzees and bonobos, gorillas spend a greater percentage of their daily time feeding and shift to herbaceous vegetation when fruits are scarce. We thus used the African apes to test the hypothesis that chewing with unusually high frequency is correlated with the expression in the jaw adductors of a high proportion of type 1 (slow, fatigue-resistant) fibers at the expense of other fiber types (the Frequent Recruitment Hypothesis). We used immunohistochemistry to determine the presence and distribution of the four major myosin heavy chain (MHC) isoforms in the anterior superficial masseter (ASM), superficial anterior temporalis, and deep anterior temporalis of four Gorilla gorilla, two Pan paniscus, and four Pan troglodytes. Serial sections were stained against slow (MHC-1/-α-cardiac) and fast (MHC-2/-M) fibers. Fibers were counted and scored for staining intensity, and fiber cross-sectional areas (CSAs) were measured and used to estimate percentage of CSA of each MHC isoform. Hybrid fibers accounted for nearly 100% of fiber types in the masseter and temporalis of all three species, resulting in three main hybrid phenotypes. As predicted, the gorilla ASM and deep anterior temporalis comprised a greater percentage of CSA of the slower, fatigue-resistant hybrid fiber type, significantly so for the ASM (p = 0.015). Finally, the results suggest that fiber phenotype of the chewing muscles contributes to behavioral flexibility in ways that would go undetected in paleontological studies relying solely on morphology of the bony masticatory apparatus.

Premasticated food transfer by wild chimpanzee mothers with their infants: Effects of maternal parity, infant age and sex, and food properties.

Premasticated food transfer, when an individual partially breaks down food through chewing and feeds it to another individual, usually mouth-to-mouth, is described widely across human cultures. This behavior plays an important role in modern humans' strategy of complementary feeding, which involves supplementing maternal milk in infant diets with processed, easily digestible versions of adult foods. The extent to which other primates engage in premasticated food transfer with infants is unclear, as premasticated food transfers have been only occasionally reported in other ape species. We investigated premasticated food transfers in 62 mother-infant pairs of wild chimpanzees at Ngogo, Uganda, as well as unresisted food taking, when mothers passively allow infants to seize food. We evaluated the presence or absence and rates of premasticated food transfer and unresisted food taking relative to maternal parity and infant age and sex and assessed the food species and part used. We found that chimpanzee mothers regularly shared premasticated food with their infants aged between 6 months and 4 years, but they were more likely to share, and more frequently shared, with younger infants. The frequency with which females shared premasticated food may relate to maternal experience, as multiparous females shared premasticated food more often than did first-time mothers, which we did not find with unresisted food taking. Both easy-to-chew, commonly eaten foods and tougher, rarely eaten foods were shared. Premasticated food transfer and unresisted food taking may be infant-rearing strategies to facilitate the transition from a diet of exclusive maternal milk to solid food during early infancy. Premasticated food transfer in particular may provide energetic, immune, or growth benefits to infants through reduced chewing effort and maternal saliva. Given our findings in chimpanzees and earlier reports in other ape species, we suggest that the foundation of complementary feeding, a uniquely hominin strategy, might have been present in a common ancestor shared with the other great apes in the form of premasticated, mouth-to-mouth food transfer by mothers with their offspring.

Age at first molar emergence in Pan troglodytes verus and variation in the timing of molar emergence among free-living chimpanzees.

Age at lower first molar (M1) emergence is a commonly used proxy for inferring life-history scheduling in fossil primates, but its utility is dependent on knowing to what extent extant populations vary in this datum and how this variation correlates with the scheduling of life-history variables. Here, we address the first of these issues among extant chimpanzees. While age at M1 emergence has been documented in several live individuals from the Kanyawara population of Pan troglodytes schweinfurthii in Uganda, it has been estimated for only one individual of Pan troglodytes verus, based on a deceased animal from the Taï Forest in Côte d'Ivoire. To further explore interpopulation variation in this variable in chimpanzees, and using dental histology, we calculated ages at death for two wild-shot individuals of P. t. verus with erupting M1, both collected in Liberia during the mid-1950s, and estimated ages at M1 emergence from the ages at death. The overall range for these two individuals is ∼4.2-4.6 yr, compared with an age of ∼3.7 yr for the individual from the Taï Forest, and <2.5-3.3 yr for the several individuals of P. t. schweinfurthii. While the absolute range of ∼2 yr in these samples combined is little greater than in captive chimpanzees, the disparity between the samples of P. t. schweinfurthii and P. t. verus is striking, although it cannot be determined if this disparity represents a subspecies difference or simply population differences expressed in two different subspecies. While life-history data are unavailable for the population to which the Liberian chimpanzees belonged, the difference in M1 emergence ages between these individuals and those from Kanyawara still suggests caution when attempting even broad life-history inference in fossil apes and hominins based on age at M1 emergence.

Root growth and dental eruption in modern human deciduous teeth with preliminary observations on great apes.

Recent studies of dental development have indicated that root growth rates are linked to the eruption of some permanent tooth types in modern humans and Pan troglodytes. Little is known about the potential links between these aspects of dental development in deciduous teeth of any primate species. This histology study calculates the rate at which roots extend in length for human deciduous maxillary teeth and a small sample of deciduous canines and premolars from P. troglodytes and Pongo pygmaeus. Links are sought between root extension rates and previously published data for deciduous tooth emergence in each of these species. Results reported here provide the first evidence that the roots of human deciduous incisors, canines, and premolars extend in length at an accelerated rate as these teeth emerge. Accelerated extension rates in a deciduous canine from Pan coincided with the age that this tooth type emerged in captive chimpanzees. High extension rates in a canine from Pongo preceded emergence age. Preliminary observations indicate that deciduous canine and premolar roots of Pan and Pongo extend in length rapidly when compared to these tooth types from modern human children. This study provides a starting point from which to investigate new links between the incremental development of deciduous roots and tooth emergence in primates.

Dust affects chewing efficiency and tooth wear in forest dwelling Western chimpanzees (Pan troglodytes verus).

OBJECTIVES: In humans it has been shown that abrasive particles in the diet result in increased tooth wear and less intense chewing behavior, both of which decrease chewing efficiency. This behavioral response may also exist in non-human primates as a means to reduce the wear effect of dust-laden food. Here we tested whether the periodical occurrence of abrasive dust particles in the diet of Western chimpanzees affects tooth wear and reduces chewing efficiency. MATERIALS AND METHODS: We measured fecal particle size of undigested food matter as an indicator of chewing efficiency in 13 Western chimpanzees of the Taï National Park (Ivory Coast) before (wet), after (wet) and during a dust-rich (dry) period. Moreover, feeding data were compiled for a further 12 chimpanzees and matched to three-dimensional surface texture data measured on two molar facets of 26 skulls of the same population. RESULTS: Fecal particles were larger during the dry period, indicating a reduced chewing efficiency compared to wet periods; age and sex did not have an effect. Concomitantly, dust led to an increase of abrasive wear evidenced by smaller texture features and higher density of fine furrows on wear facets. DISCUSSION: Our findings show that a periodical increase in dust loads on foods places a dietary-physiological stress on the digestive system in chimpanzees. We suggest that the impact of extrinsic abrasive particles from globally acting periodical dust-laden winds may affect evolutionary fitness. Further studies are required to elucidate this relationship in other non-human primates and fossil hominins.

Evaluating the triplet hypothesis during rhythmic mastication in primates.

Mammalian mastication involves precise jaw movements including transverse movement of the mandible during the power stroke. Jaw elevation and transverse movement are driven by asymmetrical jaw elevator muscle activity, which is thought to include a phylogenetically primitive and conserved triplet motor pattern consisting of: triplet I (balancing side: superficial masseter and medial pterygoid; working side: posterior temporalis), which reaches onset, peak and offset first; and triplet II (working side: superficial masseter and medial pterygoid; balancing side: posterior temporalis), which is active second. Although the presence of a triplet motor pattern has been confirmed in several primate species, the prevalence of this motor pattern - i.e. the proportion of masticatory cycles that display it - has not been evaluated in primates. The present study quantifies the presence and prevalence of the triplet motor pattern in five different primate species, Eulemur fulvus, Propithecus verreauxi, Papio anubis, Macacafuscata and Pan troglodytes, using mean onset, peak and offset time relative to working superficial masseter. In all five of the species studied, the mean triplet motor pattern was observed at peak muscle activation, and in four out of the five species the triplet motor pattern occurred more frequently than expected at random at peak muscle activation and offset. Non-triplet motor patterns were observed in varying proportions at different time points in the masticatory cycle, suggesting that the presence or absence of the triplet motor pattern is not a binomial trait. Instead, the primate masticatory motor pattern is malleable within individual cycles, within individual animals and therefore within species.

The presence of females induces elevated cortisol levels in an alpha male: Experimental evidence in chimpanzees.

OBJECTIVES: In group-living primates, it has been reported that the alpha male exhibits high concentrations of cortisol and testosterone in the context of mating competition. We investigated how the presence of females affected salivary cortisol and testosterone levels in males from a small captive group of chimpanzees (Pan troglodytes). Specifically, we assessed whether the presence of females resulted in a rapid increase in salivary cortisol and testosterone levels in the alpha male. MATERIALS AND METHODS: We compared the social behavior and salivary hormone concentrations of four males before and after the presentation of receptive females. Three times a day, we collected saliva samples, a useful matrix for investigating short-term hormonal changes, and measured cortisol and testosterone concentration by liquid chromatography-tandem mass spectrometry (LC-MS/MS). RESULTS: The frequency of inter-male aggression increased in the presence of females, indicating intense competition among males. Salivary cortisol levels increased in all males in the presence of females; however, the increase was significantly more pronounced in the alpha male. We found a complex three-way interaction among the presence of females, sampling timings, and male dominance rank in the analysis of salivary testosterone. Contrary to our prediction, a post hoc analysis revealed that salivary testosterone levels decreased after female introduction and that the alpha male did not show a higher level of salivary testosterone. CONCLUSIONS: Our study provides experimental evidence suggesting that the presence of females plays a significant role in the rank-related variation in the cortisol levels in male chimpanzees. Furthermore, our findings demonstrate the usefulness of salivary hormones for detecting short-term physiological changes in studies of socioendocrinology.

Neither chimpanzee nor human, Ardipithecus reveals the surprising ancestry of both.

Australopithecus fossils were regularly interpreted during the late 20th century in a framework that used living African apes, especially chimpanzees, as proxies for the immediate ancestors of the human clade. Such projection is now largely nullified by the discovery of Ardipithecus. In the context of accumulating evidence from genetics, developmental biology, anatomy, ecology, biogeography, and geology, Ardipithecus alters perspectives on how our earliest hominid ancestors--and our closest living relatives--evolved.

First molar eruption, weaning, and life history in living wild chimpanzees.

Understanding dental development in chimpanzees, our closest living relatives, is of fundamental importance for reconstructing the evolution of human development. Most early hominin species are believed to show rapid ape-like patterns of development, implying that a prolonged modern human childhood evolved quite recently. However, chimpanzee developmental standards are uncertain because they have never been based on living wild individuals. Furthermore, although it is well established that first molar tooth emergence (movement into the mouth) is correlated with the scheduling of growth and reproduction across primates broadly, its precise relation to solid food consumption, nursing behavior, or maternal life history is unknown. To address these concerns we conducted a photographic study of subadult chimpanzees (Pan troglodytes schweinfurthii) in Kanyawara, Kibale National Park, Uganda. Five healthy infants emerged their lower first molars (M1s) by or before 3.3 y of age, nearly identical to captive chimpanzee mean ages (∼3.2 y, n = 53). First molar emergence in these chimpanzees does not directly or consistently predict the introduction of solid foods, resumption of maternal estrous cycling, cessation of nursing, or maternal interbirth intervals. Kanyawara chimpanzees showed adult patterns of solid food consumption by the time M1 reached functional occlusion, spent a greater amount of time on the nipple while M1 was erupting than in the preceding year, and continued to suckle during the following year. Estimates of M1 emergence age in australopiths are remarkably similar to the Kanyawara chimpanzees, and recent reconstructions of their life histories should be reconsidered in light of these findings.

Dental eruption in East African wild chimpanzees.

Knowledge of chimpanzee development has played an essential role in our understanding of the evolution of human ontogeny. However, recent studies of wild ape dentitions have cast doubt on the use of developmental standards derived from captive individuals. Others have called into question the use of deceased wild individuals to infer normative development. We conducted a high resolution photographic study of living known-age subadults in the Kanyawara community (Kibale National Park, Uganda) to generate a comprehensive three year record of dental eruption (including tooth emergence ages). These non-invasive data allow comparisons of captive and wild chimpanzees, establish accurate developmental standards for relatively healthy wild individuals, and facilitate direct assessments of primate-wide associations between dental development and life history. Emergence ages in the Kanyawara chimpanzees are very similar to living Gombe chimpanzees, and are broadly comparable to deceased Taï Forest chimpanzees. Early-emerging teeth such as the deciduous dentition and first molar (M1) appear during a time of maternal dependence, and are almost indistinguishable from captive chimpanzee emergence ages, while later forming teeth in the Kanyawara population emerge in the latter half of captive age ranges or beyond. Five juveniles whose lower M1s emerged by or before 3.3 years of age continued to nurse for a year or more beyond M1 emergence, and their mothers showed considerable variation in reproductive rates. The third molars of two adolescent females emerged several months to several years prior to the birth of their first offspring. Given that broad primate-wide relationships between molar emergence and life history do not necessarily hold within this population of chimpanzees, particularly for variables that are reported to be coincident with molar emergence, we suggest that further study is required in order to predict life history variables in hominins or hominoids.

A multivariate approach to assess variation in tooth mineralization using free-lived and captive-raised chimpanzees (P. troglodytes).

OBJECTIVES: Understanding variation in dental development among primates is important to accurately characterize species-specific sequences and times of tooth formation. Conventional approaches that summarize dental development data (i.e., dental maturity score, DMS) inherently omit information about the full range of variation in raw scores; thus, classic bivariate analyses are limited for exploring patterns of variation in detail. Here we report a new multivariate approach to simultaneously assess all raw dental scores, for all teeth, among all individuals for all groups, thus retaining much greater detail about population-specific patterns of variation. METHODS: We scored (0-12) permanent tooth mineralization using radiographs of mandibles of captive-raised known-age chimpanzees (Pan troglodytes; n = 114) and free-lived age-unknown conspecifics (n = 54). As a test of our method we also scored free-lived baboons (Papio anubis, n = 50) because of well-described contrasting patterns of permanent molar initiation between Papio and Pan. Using principal component analysis (PCA), we investigated how crypt, crown and root formation scores covaried with each other in all three genera, and with chronological age in captive chimpanzees. RESULTS: PCA successfully captured additional detail about variation among raw scores. Also, compared to DMS, PC1 scores correlated equivalently well with known ages but had lower prediction error. CONCLUSIONS: We found different patterns of variation in scores between younger juvenile free and captive chimpanzees but saw no wholesale differences between groups. Pan and Papio showed different patterns of variation, further validating this multivariate approach to visualize, quantify and compare raw dental score datasets among primate species.

"Missing perikymata"--fact or fiction? A study on chimpanzee (Pan troglodytes verus) canines.

Recently, a lower than expected number of perikymata between repetitive furrow-type hypoplastic defects has been reported in chimpanzee canines from the Fongoli site, Senegal (Skinner and Pruetz: Am J Phys Anthropol 149 (2012) 468-482). Based on an observation in a localized enamel fracture surface of a canine of a chimpanzee from the Taï Forest (Ivory Coast), these authors inferred that a nonemergence of striae of Retzius could be the cause for the "missing perikymata" phenomenon in the Fongoli chimpanzees. To check this inference, we analyzed the structure of outer enamel in three chimpanzee canines. The teeth were studied using light-microscopic and scanning-electron microscopic techniques. Our analysis of the specimen upon which Skinner and Pruetz (Am J Phys Anthropol 149 (2012) 468-482) had made their original observation does not support their hypothesis. We demonstrate that the enamel morphology described by them is not caused by a nonemergence of striae of Retzius but can be attributed to structural variations in outer enamel that result in a differential fracture behavior. Although rejecting the presumed existence of nonemergent striae of Retzius, our study provided evidence that, in furrow-type hypoplastic defects, a pronounced tapering of Retzius increments can occur, with the striae of Retzius forming acute angles with the outer enamel surface. We suggest that in such cases the outcrop of some striae of Retzius is essentially unobservable at the enamel surface, causing too low perikymata counts. The pronounced tapering of Retzius increments in outer enamel presumably reflects a mild to moderate disturbance of the function of late secretory ameloblasts.

Grips and hand movements of chimpanzees during feeding in Mahale Mountains National Park, Tanzania.

It has long been assumed that stone tool making was a major factor in the evolution of derived hominin hand morphology. However, stresses on the hand associated with food retrieval and processing also have been recognized as relevant early hominin behaviors that should be investigated. To this end, chimpanzee food manipulation was videotaped in the Mahale Mountains National Park, Tanzania. Grips and hand movements by 39 chimpanzees were analyzed for arboreal and terrestrial feeding involving 10 food-types and associated vegetation. It was predicted that (1) new grips would be found that had not been observed in captivity, (2) forceful precision grips would be absent from the repertoire, as in captivity, and (3) precision handling would be observed. New grips involving the full thumb and buttressed index finger, and a new integrated pattern of grips and forceful hand movements were discovered, associated with feeding on large fruits and meat. Participation of the full thumb in these grips, rather than the distal thumb and fingers, throws light on feeding behaviors that may have become increasingly significant factors in the evolution of derived hominin thumb morphology. The proximal thumb stabilizes food with the flexed index finger against the pull of the teeth and provides leverage in breaking food into portions. Isolated qualitative observations of possibly forceful pinch by the thumb and side of the index finger highlight the need for comparative quantitative data to test whether humans are unique in forceful precision gripping capability. Precision handling was not seen.

Niche differentiation and dietary seasonality among sympatric gorillas and chimpanzees in Loango National Park (Gabon) revealed by stable isotope analysis.

The feeding ecology of sympatric great ape species yields valuable information for palaeodietary reconstructions in sympatric early hominin species. However, no isotopic references on sympatrically living apes and their feeding ecology are currently available. Here we present the first isotopic study on sympatric great apes, namely western lowland gorillas (Gorilla gorilla gorilla) and central chimpanzees (Pan troglodytes troglodytes) from Loango National Park, Gabon. We successfully analyzed the stable carbon and nitrogen isotope ratios in a selection of food plants (n = 31) and hair samples (n = 30) retrieved from sleeping nests to test whether niche partitioning among sympatric chimpanzees and gorillas is detectable using isotope analysis of hair. Ape hair strands with roots were sectioned into sequential segments (n = 100) to investigate temporal isotopic variation related to seasonal variations in food resources. We found significant δ(13)C differences between herbaceous plants and fruits, most likely due to canopy effects. While the δ(13)C values of chimpanzees indicate the consumption of fruit, the low δ(13)C values in gorilla hair indicate folivory, most likely the consumption of (13)C-depleted herbaceous vegetation. Our isotopic data also confirmed dietary overlap between chimpanzees and gorillas, which varied by season. Gorillas showed significant variation in δ(13)C values in response to season due to shifting proportions of herbaceous plants versus fruits. In chimpanzees, significant seasonal variation in δ(15)N was likely related to the seasonal availability of fruit species with particularly high δ(15)N values. In summary, we found isotopic evidence for niche partitioning and seasonal dietary variation among sympatric great apes at Loango. These findings provide a valuable reference for palaeodietary research on fossil hominins using δ(13)C analyses, particularly for studies focusing on sympatric taxa and on temporal isotopic variation within incremental tissues such as tooth enamel.

Stable nitrogen isotope analysis of dentine serial sections elucidate sex differences in weaning patterns of wild chimpanzees (Pan troglodytes).

Offspring provisioning is one of the most energetically demanding aspects of reproduction for female mammals. Variation in lactation length and weaning strategies between chimpanzees (Pan troglodytes), our closest living relative, and modern human societies have been reported. When and why these changes occurred is frequently debated. Our study used stable nitrogen isotope data of tooth root dentine from wild Western chimpanzees (Pan troglodytes verus) in Taï National Park, Côte d'Ivoire, to quantify weaning in these chimpanzees and explore if infant sex plays a role in maternal investment. We analyzed serial sections of deciduous lateral incisor root dentine from four Taï chimpanzees to establish the δ(15) N signal of nursing infants; we then analyzed serial sections of first permanent mandibular molar root dentine from 12 Taï chimpanzees to provide quantitative δ(15) N data on weaning in this population. Up to 2 years of age both sexes exhibited dentine δ(15) N values ≈2-3‰ higher than adult female Taï chimpanzees, consistent with a nursing signal. Thereafter a steady decrease in δ(15) N values consistent with the onset, and progression, of weaning, was visible. Sex differences were also evident, where male δ(15) N values decreased at a significantly slower rate compared to females. Confirmation of sex differences in maternal investment among Taï chimpanzees, demonstrates the viability of using isotope analysis to investigate weaning in non-human primates. Additionally, assuming that behaviors observed in the Taï chimpanzees are illustrative of the ancestral pattern, our results provide a platform to enable the trajectory of weaning in human evolution to be further explored.

Chimpanzee utterances refute purported missing links for novel vocalizations and syllabic speech.

Nonhuman great apes have been claimed to be unable to learn human words due to a lack of the necessary neural circuitry. We recovered original footage of two enculturated chimpanzees uttering the word "mama" and subjected recordings to phonetic analysis. Our analyses demonstrate that chimpanzees are capable of syllabic production, achieving consonant-to-vowel phonetic contrasts via the simultaneous recruitment and coupling of voice, jaw and lips. In an online experiment, human listeners naive to the recordings' origins reliably perceived chimpanzee utterances as syllabic utterances, primarily as "ma-ma", among foil syllables. Our findings demonstrate that in the absence of direct data-driven examination, great ape vocal production capacities have been underestimated. Chimpanzees possess the neural building blocks necessary for speech.

Chimpanzee fauna isotopes provide new interpretations of fossil ape and hominin ecologies.

Carbon and oxygen stable isotopes within modern and fossil tooth enamel record the aspects of an animal's diet and habitat use. This investigation reports the first isotopic analyses of enamel from a large chimpanzee community and associated fauna, thus providing a means of comparing fossil ape and early hominin palaeoecologies with those of a modern ape. Within Kibale National Park forest, oxygen isotopes differentiate primate niches, allowing for the first isotopic reconstructions of degree of frugivory versus folivory as well as use of arboreal versus terrestrial resources. In a comparison of modern and fossil community isotopic profiles, results indicate that Sivapithecus, a Miocene ape from Pakistan, fed in the forest canopy, as do chimpanzees, but inhabited a forest with less continuous canopy or fed more on leaves. Ardipithecus, an early hominin from Ethiopia, fed both arboreally and terrestrially in a more open habitat than inhabited by chimpanzees.

Diurnal variation in nutrients and chimpanzee foraging behavior.

Primate feeding behavior varies over long (e.g., weekly, seasonally, yearly) and short (e.g., hourly) scales of time due to changes in resource availability and the nutritional composition of foods. While the factors that affect long-term changes in feeding behavior have received considerable attention, few data exist regarding what drives variability in feeding behavior over the course of a single day. To address this problem, we investigated diurnal variation in chimpanzee feeding on the leaves of two species of saplings, Pterygota mildbraedii and Celtis africana, at Ngogo, Kibale National Park, Uganda. Specifically, we related short-term changes in chimpanzee feeding behavior on these leaves to diurnal variation in their nutritional composition. Results showed that chimpanzees fed on the leaves of both saplings more in the evening than they did in the morning. The nutritional quality of leaves also improved over the course of the day. Concentrations of cellulose and lignin were lower and total nonstructural carbohydrates (including sugars and starch) were higher in the evening for P. mildbraedii, and sugars were higher in the evening for C. africana. These data suggest that chimpanzees consume these resources when their quality is highest, and consequently, may track the nutrient composition of their foods over very short periods that span only a few hours. In the future, foods collected for analyses must control for time of sampling to ensure biologically meaningful assays of nutrient composition.

Facial Expression of TIPI Personality and CHMP-Tri Psychopathy Traits in Chimpanzees (Pan troglodytes) : Evidence for Honest Signalling?

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.

Demographic and hormonal evidence for menopause in wild chimpanzees.

Among mammals, post-reproductive life spans are currently documented only in humans and a few species of toothed whales. Here we show that a post-reproductive life span exists among wild chimpanzees in the Ngogo community of Kibale National Park, Uganda. Post-reproductive representation was 0.195, indicating that a female who reached adulthood could expect to live about one-fifth of her adult life in a post-reproductive state, around half as long as human hunter-gatherers. Post-reproductive females exhibited hormonal signatures of menopause, including sharply increasing gonadotropins after age 50. We discuss whether post-reproductive life spans in wild chimpanzees occur only rarely, as a short-term response to favorable ecological conditions, or instead are an evolved species-typical trait as well as the implications of these alternatives for our understanding of the evolution of post-reproductive life spans.