Flexible grouping patterns in a western and eastern chimpanzee community.

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.

DeepWild: Application of the pose estimation tool DeepLabCut for behaviour tracking in wild chimpanzees and bonobos.

Studying animal behaviour allows us to understand how different species and individuals navigate their physical and social worlds. Video coding of behaviour is considered a gold standard: allowing researchers to extract rich nuanced behavioural datasets, validate their reliability, and for research to be replicated. However, in practice, videos are only useful if data can be efficiently extracted. Manually locating relevant footage in 10,000 s of hours is extremely time-consuming, as is the manual coding of animal behaviour, which requires extensive training to achieve reliability. Machine learning approaches are used to automate the recognition of patterns within data, considerably reducing the time taken to extract data and improving reliability. However, tracking visual information to recognise nuanced behaviour is a challenging problem and, to date, the tracking and pose-estimation tools used to detect behaviour are typically applied where the visual environment is highly controlled. Animal behaviour researchers are interested in applying these tools to the study of wild animals, but it is not clear to what extent doing so is currently possible, or which tools are most suited to particular problems. To address this gap in knowledge, we describe the new tools available in this rapidly evolving landscape, suggest guidance for tool selection, provide a worked demonstration of the use of machine learning to track movement in video data of wild apes, and make our base models available for use. We use a pose-estimation tool, DeepLabCut, to demonstrate successful training of two pilot models of an extremely challenging pose estimate and tracking problem: multi-animal wild forest-living chimpanzees and bonobos across behavioural contexts from hand-held video footage. With DeepWild we show that, without requiring specific expertise in machine learning, pose estimation and movement tracking of free-living wild primates in visually complex environments is an attainable goal for behavioural researchers.

L'étude du comportement animal nous permet de comprendre comment différentes espèces et différents individus naviguent dans leur monde physique et social. Le codage vidéo du comportement est considéré comme une référence: il permet aux chercheurs d'extraire des ensembles de données comportementales riches et nuancées, de valider leur fiabilité et de reproduire les recherches. Toutefois, dans la pratique, les vidéos ne sont utiles que si les données peuvent être extraites efficacement. La localisation manuelle de séquences pertinentes parmi des dizaines de milliers d'heures prend énormément de temps, tout comme le codage manuel du comportement animal, qui nécessite une formation approfondie pour être fiable. Les approches d'apprentissage automatique sont utilisées pour automatiser la reconnaissance de modèles dans les données, ce qui réduit considérablement le temps nécessaire à l'extraction des données et améliore la fiabilité. Toutefois, le suivi des informations visuelles pour reconnaître un comportement nuancé est un problème difficile et, à ce jour, les outils de suivi et d'estimation de la pose utilisés pour détecter le comportement sont généralement appliqués lorsque l'environnement visuel est hautement contrôlé. Les chercheurs en comportement animal sont intéressés par l'application de ces outils à l'étude des animaux sauvages, mais il n'est pas clair dans quelle mesure cela est actuellement possible, ni quels outils sont les mieux adaptés à des problèmes particuliers. Pour combler ce manque de connaissances, nous décrivons les nouveaux outils disponibles dans ce paysage en évolution rapide, proposons des conseils pour la sélection des outils, fournissons une démonstration pratique de l'utilisation de l'apprentissage automatique pour suivre les mouvements dans les données vidéo des grands singes sauvages et mettons nos modèles de base à disposition pour utilisation. Nous utilisons un outil d'estimation de la pose, DeepLabCut, pour démontrer l'apprentissage réussi de deux modèles pilotes d'un problème extrêmement difficile d'estimation et de suivi de la pose: les chimpanzés et les bonobos sauvages vivant dans la forêt et représentant plusieurs animaux dans différents contextes comportementaux à partir de séquences vidéo tenues à la main. Avec DeepWild, nous montrons que, sans nécessiter d'expertise spécifique en apprentissage automatique, l'estimation de la pose et le suivi des mouvements de primates sauvages vivant en liberté dans des environnements visuellement complexes est un objectif réalisable pour les chercheurs en comportement.

To drum or not to drum: Selectivity in tree buttress drumming by chimpanzees (Pan troglodytes verus) in the Nimba Mountains, Guinea.

Chimpanzees live in fission-fusion social organizations, which means that party size, composition, and spatial distribution are constantly in flux. Moreover, chimpanzees use a remarkably extensive repertoire of vocal and nonvocal forms of communication, thought to help convey information in such a socially and spatially dynamic setting. One proposed form of nonvocal communication in chimpanzees is buttress drumming, in which an individual hits a tree buttress with its hands and/or feet, thereby producing a low-frequency acoustic signal. It is often presumed that this behavior functions to communicate over long distances and is, therefore, goal-oriented. If so, we would expect chimpanzees to exhibit selectivity in the choice of trees and buttresses used in buttress drumming. Selectivity is a key attribute of many other goal-directed chimpanzee behaviors, such as nut-cracking and ant dipping. Here, we investigate whether chimpanzees at the Seringbara study site in the Nimba Mountains, Guinea, West Africa, show selectivity in their buttress drumming behavior. Our results indicate that Seringbara chimpanzees are more likely to use larger trees and select buttresses that are thinner and have a greater surface area. These findings imply that tree buttress drumming is not a random act, but rather goal-oriented and requires knowledge of suitable trees and buttresses. Our results also point to long-distance communication as a probable function of buttress drumming based on selectivity for buttress characteristics likely to impact sound propagation. This study provides a foundation for further assessing the cognitive underpinnings and functions of buttress drumming in wild chimpanzees.

Joint Attention in Human and Chimpanzee Infants in Varied Socio-Ecological Contexts.

Joint attention (JA) is an early manifestation of social cognition, commonly described as interactions in which an infant looks or gestures to an adult female to share attention about an object, within a positive emotional atmosphere. We label this description the JA phenotype. We argue that characterizing JA in this way reflects unexamined assumptions which are, in part, due to past developmental researchers' primary focus on western, middle-class infants and families. We describe a range of cultural variations in caregiving practices, socialization goals, and parenting ethnotheories as an essential initial step in viewing joint attention within inclusive and contextualized perspectives. We begin the process of conducting a decolonized study of JA by considering the core construct of joint attention (i.e., triadic connectedness) and adopting culturally inclusive definitions (labeled joint engagement [JE]). Our JE definitions allow for attention and engagement to be expressed in visual and tactile modalities (e.g., for infants experiencing distal or proximal caregiving), with various social partners (e.g., peers, older siblings, mothers), with a range of shared topics (e.g., representing diverse socialization goals, and socio-ecologies with and without toys), and with a range of emotional tone (e.g., for infants living in cultures valuing calmness and low arousal, and those valuing exuberance). Our definition of JE includes initiations from either partner (to include priorities for adult-led or child-led interactions). Our next foundational step is making an ecological commitment to naturalistic observations (Dahl, 2017, Child Dev Perspect, 11(2), 79-84): We measure JE while infants interact within their own physical and social ecologies. This commitment allows us to describe JE as it occurs in everyday contexts, without constraints imposed by researchers. Next, we sample multiple groups of infants drawn from diverse socio-ecological settings. Moreover, we include diverse samples of chimpanzee infants to compare with diverse samples of human infants, to investigate the extent to which JE is unique to humans, and to document diversity both within and between species. We sampled human infants living in three diverse settings. U.K. infants (n = 8) were from western, middle-class families living near universities in the south of England. Nso infants (n = 12) were from communities of subsistence farmers in Cameroon, Africa. Aka infants (n = 10) were from foraging communities in the tropical rain forests of Central African Republic, Africa. We coded behavioral details of JE from videotaped observations (taken between 2004 and 2010). JE occurred in the majority of coded intervals (Mdn = 68%), supporting a conclusion that JE is normative for human infants. The JA phenotype, in contrast, was infrequent, and significantly more common in the U.K. (Mdn = 10%) than the other groups (Mdn < 3%). We found significant within-species diversity in JE phenotypes (i.e., configurations of predominant forms of JE characteristics). We conclude that triadic connectedness is very common in human infants, but there is significant contextualization of behavioral forms of JE. We also studied chimpanzee infants living in diverse socio-ecologies. The PRI/Zoo chimpanzee infants (n = 7) were from captive, stable groups of mixed ages and sexes, and included 4 infants from the Chester Zoo, U.K. and 3 from the Primate Research Institute, Kyoto University, Japan. The Gombe chimpanzee infants (n = 12) were living in a dynamically changing, wild community in the Gombe National Park, Tanzania, Africa. Additionally, we include two Home chimpanzee infants who were reared from birth by a female scientist, in the combined U.S., middle-class contexts of home and university cognition laboratory. JE was coded from videotaped observations (taken between 1993 and 2006). JE occurred during the majority of coded intervals (Mdn = 64%), consistent with the position that JE is normative for chimpanzee infants. The JA phenotype, in contrast, was rare, but more commonly observed in the two Home chimpanzee infants (in 8% and 2% of intervals) than in other chimpanzee groups (Mdns = 0%). We found within-species diversity in the configurations comprising the JE phenotypes. We conclude that triadic connectedness is very common in chimpanzee infants, but behavioral forms of joint engagement are contextualized. We compared JE across species, and found no species-uniqueness in behavioral forms, JE characteristics, or JE phenotypes. Both human and chimpanzee infants develop contextualized social cognition. Within-species diversity is embraced when triadic connectedness is described with culturally inclusive definitions. In contrast, restricting definitions to the JA phenotype privileges a behavioral form most valued in western, middle-class socio-ecologies, irrespective of whether the interactions involve human or chimpanzee infants. Our study presents a model for how to decolonize an important topic in developmental psychology. Decolonization is accomplished by defining the phenomenon inclusively, embracing diversity in sampling, challenging claims of human-uniqueness, and having an ecological commitment to observe infant social cognition as it occurs within everyday socio-ecological contexts. It is essential that evolutionary and developmental theories of social cognition are re-built on more inclusive and decolonized empirical foundations.

Body Site and Body Orientation Preferences during Social Grooming: A Comparison between Wild and Captive Chimpanzees and Bonobos.

Grooming site preferences have been relatively well studied in monkey species in order to investigate the function of social grooming. They are not only influenced by the amount of ectoparasites, but also by different social variables such as the dominance rank between individuals or their levels of affiliation. However, studies on this topic mainly come from monkey species, with almost no report on great apes. This study aimed to explore whether body site and body orientation preferences during social grooming show species-specific differences (bonobos vs. chimpanzees) and environment-specific differences (captivity vs. wild). Results showed that bonobos groomed the head, the front and faced each other more often than chimpanzees, while chimpanzees groomed the back, anogenitals and more frequently in face-to-back positions. Moreover, captive individuals were found to groom facing one another more often than wild ones, whereas wild individuals groomed the back and in face-to-back positions more. While future studies should expand their scope to include more populations per condition, our preliminary 2 by 2 comparison study highlights the influence of (i) species-specific social differences such as social tolerance, social attention and facial communication, and (ii) socioenvironmental constraints such as risk of predation, spatial crowding and levels of hygiene, that might be the two important factors determining the grooming patterns in two Panspecies.

Lethal aggression in Pan is better explained by adaptive strategies than human impacts.

Observations of chimpanzees (Pan troglodytes) and bonobos (Pan paniscus) provide valuable comparative data for understanding the significance of conspecific killing. Two kinds of hypothesis have been proposed. Lethal violence is sometimes concluded to be the result of adaptive strategies, such that killers ultimately gain fitness benefits by increasing their access to resources such as food or mates. Alternatively, it could be a non-adaptive result of human impacts, such as habitat change or food provisioning. To discriminate between these hypotheses we compiled information from 18 chimpanzee communities and 4 bonobo communities studied over five decades. Our data include 152 killings (n = 58 observed, 41 inferred, and 53 suspected killings) by chimpanzees in 15 communities and one suspected killing by bonobos. We found that males were the most frequent attackers (92% of participants) and victims (73%); most killings (66%) involved intercommunity attacks; and attackers greatly outnumbered their victims (median 8:1 ratio). Variation in killing rates was unrelated to measures of human impacts. Our results are compatible with previously proposed adaptive explanations for killing by chimpanzees, whereas the human impact hypothesis is not supported.

The macronutrient composition of wild and cultivated plant foods of West African chimpanzees (Pan troglodytes verus) inhabiting an anthropogenic landscape.

Agricultural expansion encroaches on tropical forests and primates in such landscapes frequently incorporate crops into their diet. Understanding the nutritional drivers behind crop-foraging can help inform conservation efforts to improve human-primate coexistence. This study builds on existing knowledge of primate diets in anthropogenic landscapes by estimating the macronutrient content of 24 wild and 11 cultivated foods (90.5% of food intake) consumed by chimpanzees (Pan troglodytes verus) at Bossou, Guinea, West Africa. We also compared the macronutrient composition of Bossou crops to published macronutrient measures of crops from Bulindi, Uganda, East Africa. The composition of wild fruits, leaves, and pith were consistent with previous reports for primate diets. Cultivated fruits were higher in carbohydrates and lower in insoluble fiber than wild fruits, while wild fruits were higher in protein. Macronutrient content of cultivated pith fell within the ranges of consumed wild pith. Oil palm food parts were relatively rich in carbohydrates, protein, lipids, and/or fermentable fiber, adding support for the nutritional importance of the oil palm for West African chimpanzees. We found no differences in the composition of cultivated fruits between Bossou and Bulindi, suggesting that macronutrient content alone does not explain differences in crop selection. Our results build on the current understanding of chimpanzee feeding ecology within forest-agricultural mosaics and provide additional support for the assumption that crops offer primates energetic benefits over wild foods.

Investigating the Function of Mutual Grooming in Captive Bonobos (Pan paniscus) and Chimpanzees (Pan troglodytes).

Social grooming is often exchanged between individuals in many primate species. Rates of bidirectional (or simultaneous mutual) grooming vary across primate species, and its function is not yet fully understood. For example, mutual grooming is frequent in chimpanzees but rare in most primate species including wild bonobos. There are, however, no quantitative data available in captive bonobos. Therefore, through the direct comparison between captive bonobos and chimpanzees, this study aimed to (i) compare the frequency of mutual grooming between Pan species, (ii) explore and compare the function of mutual grooming, and (iii) discuss the rarity of this behavior in wild bonobo populations. We tested three hypotheses following the previous literature in wild chimpanzees. The social bonding hypothesis states that mutual grooming facilitates the maintenance of strong dyadic bonds. The immediate investment hypothesis states that it serves to signal willingness to invest in the interaction. The switching hypothesis states that mutual grooming serves no function but only occurs as an overlap to change the direction of unidirectional grooming. Our findings strongly supported the immediate investment hypothesis, but not the others. Grooming bouts that included mutual grooming were longer and more equitable than bouts without, illustrating that captive Pan species use mutual grooming to maximize their short-term benefits and increase the social value of their interaction. Captive bonobos performed mutual grooming in similar proportions and for similar functions as captive and wild chimpanzees do. This contrasts with wild bonobos who engage in this behavior only rarely. We suggest that the differences in patterns of mutual grooming between captive and wild bonobos might be explained by different degrees of kinship or by a potential intraspecies variation.

Crab-fishing by chimpanzees in the Nimba Mountains, Guinea.

The significance of aquatic food resources for hominins is poorly understood, despite evidence of consumption as early as 1.95 million years ago (Ma). Here we present the first evidence of a non-human ape habitually catching and consuming aquatic crabs. Chimpanzees (Pan troglodytes verus) in the rainforest of the Nimba Mountains (Guinea) consumed freshwater crabs year-round, irrespective of rainfall or ripe fruit availability. Parties of females and offspring fished for crabs more than predicted and for longer durations than adult males. Across months, crab-fishing was negatively correlated with ant-dipping, suggesting a similar nutritional role. These findings contribute to our understanding of aquatic faunivory among hominins. First, aquatic faunivory can occur in closed forests in addition to open wetlands. Second, aquatic fauna could have been a staple part of some hominin diets, rather than merely a fallback food. Third, the habitual consumption of aquatic fauna could have been especially important for females and their immature offspring. In addition to providing small amounts of essential fatty acids, crabs might also be eaten for their micronutrients such as sodium and calcium, especially by females and young individuals who may have limited access to meat.

The gesture 'Touch': Does meaning-making develop in chimpanzees' use of a very flexible gesture?

In this bottom-up study of gesture, we focused on the details of a single gesture, Touch. We compared characteristics of use by three young chimpanzees with those of 11 adults, their interactive partners, housed in a semi-natural social group at the Kyoto University Primate Research Institute (KUPRI) in Japan. Five hundred eighty-one observations of the gesture Touch were collected across a four-year time span. This single gesture had 36 different forms, was directed to 70 different target locations on the body of social partners, and occurred in 26 different contexts. Significant differences were found between infant and adult initiators in the form, target locations, and contexts of the gesture Touch. There was a wide diversity in form-location patterns within each context, and there were no form-location patterns specific to particular contexts. Thus, we demonstrate that this gesture exhibits flexibility in form and flexibility in use. The results from this study illustrate the importance of contextualized meaning in understanding flexibility in the gesture use of great apes.

Impaired Air Conditioning within the Nasal Cavity in Flat-Faced Homo.

We are flat-faced hominins with an external nose that protrudes from the face. This feature was derived in the genus Homo, along with facial flattening and reorientation to form a high nasal cavity. The nasal passage conditions the inhaled air in terms of temperature and humidity to match the conditions required in the lung, and its anatomical variation is believed to be evolutionarily sensitive to the ambient atmospheric conditions of a given habitat. In this study, we used computational fluid dynamics (CFD) with three-dimensional topology models of the nasal passage under the same simulation conditions, to investigate air-conditioning performance in humans, chimpanzees, and macaques. The CFD simulation showed a horizontal straight flow of inhaled air in chimpanzees and macaques, contrasting with the upward and curved flow in humans. The inhaled air is conditioned poorly in humans compared with nonhuman primates. Virtual modifications to the human external nose topology, in which the nasal vestibule and valve are modified to resemble those of chimpanzees, change the airflow to be horizontal, but have little influence on the air-conditioning performance in humans. These findings suggest that morphological variation of the nasal passage topology was only weakly sensitive to the ambient atmosphere conditions; rather, the high nasal cavity in humans was formed simply by evolutionary facial reorganization in the divergence of Homo from the other hominin lineages, impairing the air-conditioning performance. Even though the inhaled air is not adjusted well within the nasal cavity in humans, it can be fully conditioned subsequently in the pharyngeal cavity, which is lengthened in the flat-faced Homo. Thus, the air-conditioning faculty in the nasal passages was probably impaired in early Homo members, although they have survived successfully under the fluctuating climate of the Plio-Pleistocene, and then they moved "Out of Africa" to explore the more severe climates of Eurasia.

Chimpanzees in an anthropogenic landscape: Examining food resources across habitat types at Bossou, Guinea, West Africa.

Many primate populations occur outside protected areas in fragmented anthropogenic landscapes. Empirical data on the ecological characteristics that define an anthropogenic landscape are urgently required if conservation initiatives in such environments are to succeed. The main objective of our study was to determine the composition and availability of chimpanzee (Pan troglodytes verus) food resources across fine spatial scales in the anthropogenic landscape of Bossou, Guinea, West Africa. We examined food resources in all habitat types available in the chimpanzees' core area. We surveyed resource composition, structure and heterogeneity (20 m × 20 m quadrats, N = 54) and assessed temporal availability of food from phenology trails (total distance 5951 m; 1073 individual trees) over 1 year (2012-2013). Over half of Bossou consists of regenerating forest and is highly diverse in terms of chimpanzee food species; large fruit bearing trees are rare and confined to primary and riverine forest. Moraceae (mulberries and figs) was the dominant family, trees of which produce drupaceous fruits favored by chimpanzees. The oil palm occurs at high densities throughout and is the only species found in all habitat types except primary forest. Our data suggest that the high densities of oil palm and fig trees, along with abundant terrestrial herbaceous vegetation and cultivars, are able to provide the chimpanzees with widely available resources, compensating for the scarcity of large fruit trees. A significant difference was found between habitat types in stem density/ha and basal area m2 /ha of chimpanzee food species. Secondary, young secondary, and primary forest emerged as the most important habitat types for availability of food tree species. Our study emphasizes the importance of examining ecological characteristics of an anthropogenic landscape as each available habitat type is unlikely to be equally important in terms of spatial and temporal availability of resources. Am. J. Primatol. 78:1237-1249, 2016. © 2016 Wiley Periodicals, Inc.

A horse's eye view: size and shape discrimination compared with other mammals.

Mammals have adapted to a variety of natural environments from underwater to aerial and these different adaptations have affected their specific perceptive and cognitive abilities. This study used a computer-controlled touchscreen system to examine the visual discrimination abilities of horses, particularly regarding size and shape, and compared the results with those from chimpanzee, human and dolphin studies. Horses were able to discriminate a difference of 14% in circle size but showed worse discrimination thresholds than chimpanzees and humans; these differences cannot be explained by visual acuity. Furthermore, the present findings indicate that all species use length cues rather than area cues to discriminate size. In terms of shape discrimination, horses exhibited perceptual similarities among shapes with curvatures, vertical/horizontal lines and diagonal lines, and the relative contributions of each feature to perceptual similarity in horses differed from those for chimpanzees, humans and dolphins. Horses pay more attention to local components than to global shapes.

Primate archaeology.

All modern humans use tools to overcome limitations of our anatomy and to make difficult tasks easier. However, if tool use is such an advantage, we may ask why it is not evolved to the same degree in other species. To answer this question, we need to bring a long-term perspective to the material record of other members of our own order, the Primates.

Chimpanzees prey on army ants at Seringbara, Nimba Mountains, Guinea: predation patterns and tool use characteristics.

Chimpanzees are renowned for their use of foraging tools in harvesting social insects and some populations use tools to prey on aggressive army ants (Dorylus spp.). Tool use in army ant predation varies across chimpanzee study sites with differences in tool length, harvesting technique, and army ant species targeted. However, surprisingly little is known about the detailed ecology of army ant predation. We studied army ant predation by chimpanzees (Pan troglodytes verus) at the Seringbara study site in the Nimba Mountains, Guinea (West Africa), over 10 years (2003-2013). We investigated chimpanzee selectivity with regards to army ant prey species. We assessed the temporal variation in army ant-feeding and examined whether army ant predation was related to rainfall or ripe fruit availability. Moreover, we examined whether chimpanzees showed selectivity regarding plant species used for tool manufacture, as well as the relationship between tool species preference and tool collection distance. Lastly, we measured tool properties and investigated the use of tool sets and composite tools in army ant predation. Seringbara chimpanzees preyed on one army ant species (D. nigricans) more often than expected based on encounter rates, which may be explained by the overlap in altitudinal distribution between chimpanzees and D. nigricans. Army ant predation was not related to rainfall or fruit availability. Chimpanzees were selective in their choice of tool materials and collected their preferred tool species (Alchornea hirtella) from greater distances than they did other species. Lastly, Seringbara chimpanzees used both tool sets and composite tools (tree perch) in army ant predation. Tool types (dig vs. dip) differed in width and strength, but not length. Tool composites were found at 40% of ant-feeding sites. Our study sheds new light on the ecology of army ant predation and provides novel insights into chimpanzee selection of army ant prey and tool species.

Evidence for a midlife crisis in great apes consistent with the U-shape in human well-being.

Recently, economists and behavioral scientists have studied the pattern of human well-being over the lifespan. In dozens of countries, and for a large range of well-being measures, including happiness and mental health, well-being is high in youth, falls to a nadir in midlife, and rises again in old age. The reasons for this U-shape are still unclear. Present theories emphasize sociological and economic forces. In this study we show that a similar U-shape exists in 508 great apes (two samples of chimpanzees and one sample of orangutans) whose well-being was assessed by raters familiar with the individual apes. This U-shaped pattern or "midlife crisis" emerges with or without use of parametric methods. Our results imply that human well-being's curved shape is not uniquely human and that, although it may be partly explained by aspects of human life and society, its origins may lie partly in the biology we share with great apes. These findings have implications across scientific and social-scientific disciplines, and may help to identify ways of enhancing human and ape well-being.

The origin of representational drawing: a comparison of human children and chimpanzees.

To examine the evolutional origin of representational drawing, two experiments directly compared the drawing behavior of human children and chimpanzees. The first experiment observed free drawing after model presentation, using imitation task. From longitudinal observation of humans (N = 32, 11-31 months), the developmental process of drawing until the emergence of shape imitation was clarified. Adult chimpanzees showed the ability to trace a model, which was difficult for humans who had just started imitation. The second experiment, free drawing on incomplete facial stimuli, revealed the remarkable difference between two species. Humans (N = 57, 6-38 months) tend to complete the missing parts even with immature motor control, whereas chimpanzees never completed the missing parts and instead marked the existing parts or traced the outlines. Cognitive characteristics may affect the emergence of representational drawings.

Density estimates and nesting-site selection in chimpanzees of the Nimba Mountains, Côte d'Ivoire, and Guinea.

We investigated nesting behavior of non habituated chimpanzees populating the Nimba Mountains to document their abundance and their criterions of nesting-site selection. During a 19-month study we walked 80 km of transects and recces each month, and recorded 764 nests (mean group size = 2.23 nests) along with characteristics of vegetation structure and composition, topography, and seasonality. Population density estimated with two nest count methods ranged between 0.14 and 0.65 chimpanzee/km(2) . These values are lower than previous estimates, emphasizing the necessity of protecting remaining wild ape populations. Chimpanzees built nests in 108 tree species out of 437 identified, but 2.3% of total species comprised 52% of nests. Despite they preferred nesting in trees of 25-29 cm DBH and at a mean height of 8.02 m, we recorded an important proportion of terrestrial nests (8.2%) that may reflect a cultural trait of Nimba chimpanzees. A logistic model of nest presence formulated as a function of 12 habitat variables revealed preference for gallery and mountain forests rather than lowland forest, and old-growth forest rather than secondary forests. They nested more frequently in the study area during the dry season (December-April). The highest probability of observing nests was at 770 m altitude, particularly in steep locations (mean ground declivity = 15.54%). Several of the reported nest characteristics combined with the existence of two geographically separated clusters of nest, suggest that the study area constitutes the non-overlapping peripheral areas of two distinct communities. This nest-based study led us to findings on the behavioral ecology of Nimba chimpanzees, which constitute crucial knowledge to implement efficient and purpose-built conservation.

Visuoauditory mappings between high luminance and high pitch are shared by chimpanzees (Pan troglodytes) and humans.

Humans share implicit preferences for certain cross-sensory combinations; for example, they consistently associate higher-pitched sounds with lighter colors, smaller size, and spikier shapes. In the condition of synesthesia, people may experience such cross-modal correspondences to a perceptual degree (e.g., literally seeing sounds). So far, no study has addressed the question whether nonhuman animals share cross-modal correspondences as well. To establish the evolutionary origins of cross-modal mappings, we tested whether chimpanzees (Pan troglodytes) also associate higher pitch with higher luminance. Thirty-three humans and six chimpanzees were required to classify black and white squares according to their color while hearing irrelevant background sounds that were either high-pitched or low-pitched. Both species performed better when the background sound was congruent (high-pitched for white, low-pitched for black) than when it was incongruent (low-pitched for white, high-pitched for black). An inherent tendency to pair high pitch with high luminance hence evolved before the human lineage split from that of chimpanzees. Rather than being a culturally learned or a linguistic phenomenon, this mapping constitutes a basic feature of the primate sensory system.

The Arena System: a novel shared touch-panel apparatus for the study of chimpanzee social interaction and cognition.

We report on the development of a novel shared touch-panel apparatus for examining a diverse range of topics in great ape social cognition and interaction. Our apparatus-named the Arena System-is composed of a single multitouch monitor that spans across two separate testing booths, so that individuals situated in each booth have tactile access to half of the monitor and visual access to the whole monitor. Additional components of the system include a smart-film barrier able to restrict visual access between the booths, as well as two automated feeding devices that dispense food rewards to the subjects. The touch-panel, smart-film, and feeders are controlled by a PC that is also responsible for running the experimental tasks. We present data from a pilot behavioral game theory study with two chimpanzees in order to illustrate the efficacy of our method, and we suggest applications for a range of topics including animal social learning, coordination, and behavioral economics. The system enables fully automated experimental procedures, which means that no human participation is needed to run the tasks. The novel use of a touch-panel in a social setting allows for a finer degree of data resolution than do the traditional experimental apparatuses used in prior studies on great ape social interaction.