Many morphs: Parsing gesture signals from the noise.

Parsing signals from noise is a general problem for signallers and recipients, and for researchers studying communicative systems. Substantial efforts have been invested in comparing how other species encode information and meaning, and how signalling is structured. However, research depends on identifying and discriminating signals that represent meaningful units of analysis. Early approaches to defining signal repertoires applied top-down approaches, classifying cases into predefined signal types. Recently, more labour-intensive methods have taken a bottom-up approach describing detailed features of each signal and clustering cases based on patterns of similarity in multi-dimensional feature-space that were previously undetectable. Nevertheless, it remains essential to assess whether the resulting repertoires are composed of relevant units from the perspective of the species using them, and redefining repertoires when additional data become available. In this paper we provide a framework that takes data from the largest set of wild chimpanzee (Pan troglodytes) gestures currently available, splitting gesture types at a fine scale based on modifying features of gesture expression using latent class analysis (a model-based cluster detection algorithm for categorical variables), and then determining whether this splitting process reduces uncertainty about the goal or community of the gesture. Our method allows different features of interest to be incorporated into the splitting process, providing substantial future flexibility across, for example, species, populations, and levels of signal granularity. Doing so, we provide a powerful tool allowing researchers interested in gestural communication to establish repertoires of relevant units for subsequent analyses within and between systems of communication.

GesturalOrigins: A bottom-up framework for establishing systematic gesture data across ape species.

Current methodologies present significant hurdles to understanding patterns in the gestural communication of individuals, populations, and species. To address this issue, we present a bottom-up data collection framework for the study of gesture: GesturalOrigins. By "bottom-up", we mean that we minimise a priori structural choices, allowing researchers to define larger concepts (such as 'gesture types', 'response latencies', or 'gesture sequences') flexibly once coding is complete. Data can easily be re-organised to provide replication of, and comparison with, a wide range of datasets in published and planned analyses. We present packages, templates, and instructions for the complete data collection and coding process. We illustrate the flexibility that our methodological tool offers with worked examples of (great ape) gestural communication, demonstrating differences in the duration of action phases across distinct gesture action types and showing how species variation in the latency to respond to gestural requests may be revealed or masked by methodological choices. While GesturalOrigins is built from an ape-centred perspective, the basic framework can be adapted across a range of species and potentially to other communication systems. By making our gesture coding methods transparent and open access, we hope to enable a more direct comparison of findings across research groups, improve collaborations, and advance the field to tackle some of the long-standing questions in comparative gesture research.

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.

Towards a great ape dictionary: Inexperienced humans understand common nonhuman ape gestures.

In the comparative study of human and nonhuman communication, ape gesturing provided the first demonstrations of flexible, intentional communication outside human language. Rich repertoires of these gestures have been described in all ape species, bar one: us. Given that the majority of great ape gestural signals are shared, and their form appears biologically inherited, this creates a conundrum: Where did the ape gestures go in human communication? Here, we test human recognition and understanding of 10 of the most frequently used ape gestures. We crowdsourced data from 5,656 participants through an online game, which required them to select the meaning of chimpanzee and bonobo gestures in 20 videos. We show that humans may retain an understanding of ape gestural communication (either directly inherited or part of more general cognition), across gesture types and gesture meanings, with information on communicative context providing only a marginal improvement in success. By assessing comprehension, rather than production, we accessed part of the great ape gestural repertoire for the first time in adult humans. Cognitive access to an ancestral system of gesture appears to have been retained after our divergence from other apes, drawing deep evolutionary continuity between their communication and our own.

Dialects in leaf-clipping and other leaf-modifying gestures between neighbouring communities of East African chimpanzees.

Dialects are a cultural property of animal communication previously described in the signals of several animal species. While dialects have predominantly been described in vocal signals, chimpanzee leaf-clipping and other 'leaf-modifying' gestures, used across chimpanzee and bonobo communities, have been suggested as a candidate for cultural variation in gestural communication. Here we combine direct observation with archaeological techniques to compare the form and use of leaf-modifying gestures in two neighbouring communities of East African chimpanzees. We found that while both communities used multiple forms, primarily within sexual solicitation, they showed a strong preference for a single, different gesture form. The observed variation in form preference between these neighbouring communities within the same context suggests that these differences are, at least in part, socially derived. Our results highlight an unexplored source of variation and flexibility in gestural communication, opening the door for future research to explore socially derived dialects in non-vocal communication.

Maternal attitudes and behaviours differentially shape infant early life experience: A cross cultural study.

Early life environments afford infants a variety of learning opportunities, and caregivers play a fundamental role in shaping infant early life experience. Variation in maternal attitudes and parenting practices is likely to be greater between than within cultures. However, there is limited cross-cultural work characterising how early life environment differs across populations. We examined the early life environment of infants from two cultural contexts where attitudes towards parenting and infant development were expected to differ: in a group of 53 mother-infant dyads in the UK and 44 mother-infant dyads in Uganda. Participants were studied longitudinally from when infants were 3- to 15-months-old. Questionnaire data revealed the Ugandan mothers had more relational attitudes towards parenting than the mothers from the UK, who had more autonomous parenting attitudes. Using questionnaires and observational methods, we examined whether infant development and experience aligned with maternal attitudes. We found the Ugandan infants experienced a more relational upbringing than the UK infants, with Ugandan infants receiving more distributed caregiving, more body contact with their mothers, and more proximity to mothers at night. Ugandan infants also showed earlier physical development compared to UK infants. Contrary to our expectations, however, Ugandan infants were not in closer proximity to their mothers during the day, did not have more people in proximity or more partners for social interaction compared to UK infants. In addition, when we examined attitudes towards specific behaviours, mothers' attitudes rarely predicted infant experience in related contexts. Taken together our findings highlight the importance of measuring behaviour, rather than extrapolating expected behaviour based on attitudes alone. We found infants' early life environment varies cross-culturally in many important ways and future research should investigate the consequences of these differences for later development.

Recognition of visual kinship signals in chimpanzees (Pan troglodytes) by humans (Homo sapiens).

Associating with kin provides individual benefits but requires that these relationships be detectable. In humans, facial phenotype matching might help assess paternity; however, evidence for it is mixed. In chimpanzees, concealing visual cues of paternity may be beneficial due to their promiscuous mating system and the considerable risk of infanticide by males. On the other hand, detecting kin can also aid chimpanzees in avoiding inbreeding and in forming alliances that improve kin-mediated fitness. Although previous studies assessing relatedness based on facial resemblance in chimpanzees exist, they used images of captive populations in whom selection pressures and reproductive opportunities are controlled and only assessed maternity or paternity of adult offspring. In natural populations, the chances of infanticide are highest during early infancy, suggesting that young infants would benefit most from paternity concealment, whereas adults and subadults would benefit from the detection of all types of kin, including half-siblings. In our experiment, we conducted an online study with human participants, in which they had to assess the relatedness of chimpanzees based on facial similarity. To address previous methodological constraints, we used chimpanzee images across all ages, as well as maternal and paternal half-siblings. We found that kin status was detected above chance across all relatedness categories, with easier kin detection of father-offspring pairs, females, and older chimpanzees. Together, these findings support the existence of paternity confusion in infant chimpanzees and provide a possible mechanism for incest avoidance and kin-based social alliances in older individuals. (PsycInfo Database Record (c) 2022 APA, all rights reserved).

Are ape gestures like words? Outstanding issues in detecting similarities and differences between human language and ape gesture.

Opinion piece: ape gestures are made intentionally, inviting parallels with human language; but how similar are their gestures to words? Here we ask this in three ways, considering: flexibility and ambiguity, first- and second-order intentionality, and usage in interactive exchanges. Many gestures are used to achieve several, often very distinct, goals. Such apparent ambiguity in meaning is potentially disruptive for communication, but-as with human language-situational and interpersonal context may largely resolve the intended meaning. Our evidence for first-order intentional use of gesture is abundant, but how might we establish a case for the second-order intentional use critical to language? Finally, words are rarely used in tidy signal-response sequences but are exchanged in back-and-forth interaction. Do gestures share this property? In this paper, we examine these questions and set out ways in which they can be resolved, incorporating data from wild chimpanzees. This article is part of the theme issue 'Cognition, communication and social bonds in primates'.

A socio-ecological perspective on the gestural communication of great ape species, individuals, and social units.

Over the last 30 years, most research on non-human primate gestural communication has been produced by psychologists, which has shaped the questions asked and the methods used. These researchers have drawn on concepts from philosophy, linguistics, anthropology, and ethology, but despite these broad influences the field has neglected to situate gestures into the socio-ecological context in which the diverse species, individuals, and social-units exist. In this review, we present current knowledge about great ape gestural communication in terms of repertoires, meanings, and development. We fold this into a conversation about variation in other types of ape social behaviour to identify areas for future research on variation in gestural communication. Given the large variation in socio-ecological factors across species and social-units (and the individuals within these groups), we may expect to find different preferences for specific gesture types; different needs for communicating specific meanings; and different rates of encountering specific contexts. New tools, such as machine-learning based automated movement tracking, may allow us to uncover potential variation in the speed and form of gesture actions or parts of gesture actions. New multi-group multi-generational datasets provide the opportunity to apply analyses, such as Bayesian modelling, which allows us to examine these rich behavioural landscapes. Together, by expanding our questions and our methods, researchers may finally be able to study great ape gestures from the perspective of the apes themselves and explore what this gestural communication system reveals about apes' thinking and experience of their world.

Dominance style is a key predictor of vocal use and evolution across nonhuman primates.

Animal communication has long been thought to be subject to pressures and constraints associated with social relationships. However, our understanding of how the nature and quality of social relationships relates to the use and evolution of communication is limited by a lack of directly comparable methods across multiple levels of analysis. Here, we analysed observational data from 111 wild groups belonging to 26 non-human primate species, to test how vocal communication relates to dominance style (the strictness with which a dominance hierarchy is enforced, ranging from 'despotic' to 'tolerant'). At the individual-level, we found that dominant individuals who were more tolerant vocalized at a higher rate than their despotic counterparts. This indicates that tolerance within a relationship may place pressure on the dominant partner to communicate more during social interactions. At the species-level, however, despotic species exhibited a larger repertoire of hierarchy-related vocalizations than their tolerant counterparts. Findings suggest primate signals are used and evolve in tandem with the nature of interactions that characterize individuals' social relationships.

Detecting joint attention events in mother-infant dyads: Sharing looks cannot be reliably identified by naïve third-party observers.

Joint attention, or sharing attention with another individual about an object or event, is a critical behaviour that emerges in pre-linguistic infants and predicts later language abilities. Given its importance, it is perhaps surprising that there is no consensus on how to measure joint attention in prelinguistic infants. A rigorous definition proposed by Siposova & Carpenter (2019) requires the infant and partner to gaze alternate between an object and each other (coordination of attention) and exchange communicative signals (explicit acknowledgement of jointly sharing attention). However, Hobson and Hobson (2007) proposed that the quality of gaze between individuals is, in itself, a sufficient communicative signal that demonstrates sharing of attention. They proposed that observers can reliably distinguish "sharing", "checking", and "orienting" looks, but the empirical basis for this claim is limited as their study focussed on two raters examining looks from 11-year-old children. Here, we analysed categorisations made by 32 naïve raters of 60 infant looks to their mothers, to examine whether they could be reliably distinguished according to Hobson and Hobson's definitions. Raters had overall low agreement and only in 3 out of 26 cases did a significant majority of the raters agree with the judgement of the mother who had received the look. For the looks that raters did agree on at above chance levels, look duration and the overall communication rate of the mother were identified as cues that raters may have relied upon. In our experiment, naïve third party observers could not reliably determine the type of look infants gave to their mothers, which indicates that subjective judgements of types of look should not be used to identify mutual awareness of sharing attention in infants. Instead, we advocate the use of objective behaviour measurement to infer that interactants know they are 'jointly' attending to an object or event, and believe this will be a crucial step in understanding the ontogenetic and evolutionary origins of joint attention.

Scratching beneath the surface: intentionality in great ape signal production.

Despite important similarities having been found between human and animal communication systems, surprisingly little research effort has focussed on whether the cognitive mechanisms underpinning these behaviours are also similar. In particular, it is highly debated whether signal production is the result of reflexive processes, or can be characterized as intentional. Here, we critically evaluate the criteria that are used to identify signals produced with different degrees of intentionality, and discuss recent attempts to apply these criteria to the vocal, gestural and multimodal communicative signals of great apes and more distantly related species. Finally, we outline the necessary research tools, such as physiologically validated measures of arousal, and empirical evidence that we believe would propel this debate forward and help unravel the evolutionary origins of human intentional communication. This article is part of the theme issue 'What can animal communication teach us about human language?'

Bonobo and chimpanzee gestures overlap extensively in meaning.

Cross-species comparison of great ape gesturing has so far been limited to the physical form of gestures in the repertoire, without questioning whether gestures share the same meanings. Researchers have recently catalogued the meanings of chimpanzee gestures, but little is known about the gesture meanings of our other closest living relative, the bonobo. The bonobo gestural repertoire overlaps by approximately 90% with that of the chimpanzee, but such overlap might not extend to meanings. Here, we first determine the meanings of bonobo gestures by analysing the outcomes of gesturing that apparently satisfy the signaller. Around half of bonobo gestures have a single meaning, while half are more ambiguous. Moreover, all but 1 gesture type have distinct meanings, achieving a different distribution of intended meanings to the average distribution for all gesture types. We then employ a randomisation procedure in a novel way to test the likelihood that the observed between-species overlap in the assignment of meanings to gestures would arise by chance under a set of different constraints. We compare a matrix of the meanings of bonobo gestures with a matrix for those of chimpanzees against 10,000 randomised iterations of matrices constrained to the original data at 4 different levels. We find that the similarity between the 2 species is much greater than would be expected by chance. Bonobos and chimpanzees share not only the physical form of the gestures but also many gesture meanings.

The gestural repertoire of the wild bonobo (Pan paniscus): a mutually understood communication system.

In animal communication, signallers and recipients are typically different: each signal is given by one subset of individuals (members of the same age, sex, or social rank) and directed towards another. However, there is scope for signaller-recipient interchangeability in systems where most signals are potentially relevant to all age-sex groups, such as great ape gestural communication. In this study of wild bonobos (Pan paniscus), we aimed to discover whether their gestural communication is indeed a mutually understood communicative repertoire, in which all individuals can act as both signallers and recipients. While past studies have only examined the expressed repertoire, the set of gesture types that a signaller deploys, we also examined the understood repertoire, the set of gestures to which a recipient reacts in a way that satisfies the signaller. We found that most of the gestural repertoire was both expressed and understood by all age and sex groups, with few exceptions, suggesting that during their lifetimes all individuals may use and understand all gesture types. Indeed, as the number of overall gesture instances increased, so did the proportion of individuals estimated to both express and understand a gesture type. We compared the community repertoire of bonobos to that of chimpanzees, finding an 88 % overlap. Observed differences are consistent with sampling effects generated by the species' different social systems, and it is thus possible that the repertoire of gesture types available to Pan is determined biologically.

Long-sightedness in old wild bonobos during grooming.

Some scientists have suggested that, among Hominidae, prolonged postmenopausal longevity evolved uniquely in humans [1], while others disagree [2]. There have, however, been few empirical studies on how physiological aging and somatic durability in humans compare to our closest relatives - chimpanzees and bonobos [3]. If prolonged lifespan is selected for in humans, physiological aging, including reproductive and somatic senescence, might be different for Pan and Homo. But it seems that the parameters of reproductive senescence, such as the age of having their final offspring and the number of years between generations, are not very different between chimpanzee and human females [4]. Here, we report evidence for five cases of long-sightedness (presbyopia) in old wild bonobos, exhibited during grooming. Our results suggest that senescence of the eye has not changed much since the divergence of Pan and Homo from their common ancestor.