Catarrhine primates gesture preferentially with their right hands, which led to the hypothesis of a gestural origin of human left-hemispheric specialization for language. However, the factors influencing this gestural laterality remain understudied in non-hominoid species, particularly in intraspecific contexts, although it may bring valuable insights into the proximate and ultimate causes of language lateralization. We present here a preliminary investigation of intraspecific gestural laterality in catarrhine monkeys, red-capped mangabeys (Cercocebus torquatus). We described the spontaneous production of brachio-manual intentional gestures in twenty-five captive subjects. Although we did not evidence any significant gestural lateralization neither at the individual- nor population-level, we found that mangabeys preferentially use their right hands to gesture in negative social contexts, such as aggressions, suggesting an effect of emotional lateralization, and that they adapt to the position of their receiver by preferentially using their ipsilateral hand to communicate. These results corroborate previous findings from ape studies. By contrast, factors related to gesture form and socio-demographic characteristics of signaler and receiver did not affect gestural laterality. To understand better the relationships between gestural laterality and brain lateralization from an evolutionary perspective, we suggest that the gestural communication of other monkey species should be examined with a multifactorial approach.

Although studies on primate communication have mainly focused on single communication systems (gestural, facial or vocal), there is an increasing interest in a more integrative approach to describe the communication of our closest relatives, particularly with the aim of investigating the evolutionary roots of human language. The understanding of the ultimate functions of multimodal (involving signals of different sensory modalities) and multicomponent (association of different signal types) communication needs more systematic description of multiple signal use in primate species, with details on the circumstances leading to such complex signalling. In the present study, we describe the sequential use of communicative signals of different types and sensory modalities in a captive population of catarrhine monkeys, the red-capped mangabey, Cercocebus torquatus. We applied existing analysis methods from other disciplines to define, systematically describe and quantify the production of signals among communicative sequences. We notably used sequence analysis tools (based on dissimilarity measures) to identify the typical signal sequences produced by mangabeys, and network analysis to describe dyadic signal associations among these sequences. The focal observation of five social groups of mangabeys allowed us to identify 424 communicative sequences, which could be grouped into eight main categories, and whose complexity, multimodality and multicomponentiality depended on social context and signaller characteristics. Overall, captive mangabeys frequently associated communication signals of all types (body, facial and vocal signals) and modalities (visual, audible and tactile), in a flexible way. Our results complete previous description of red-capped mangabey signalling and highlight the need for a multimodal and multicomponent approach to understand the complexity of primate communication. Moreover, we propose the method we used as a way to enhance primate communication analysis, in the frame of comparative research. (c) 2021 The Association for the Study of Animal Behaviour. Published by Elsevier Ltd. All rights reserved.

Primate communication relies strongly on the visual modality, notably through the production of a wide range of expressive facial signals. We investigated here the facial display repertoire of a relatively little-studied cercopithecid species, red-capped mangabeys (Cercocebus torquatus), and questioned whether their facial displays were dependent on social contexts and accompanied by indices of intentionality. Although the dual intentional and emotional use of apes' facial expressions has recently been suggested, the question of whether monkeys produce facial expressions intentionally to communicate remains open. We described 6 facial displays produced by captive red-capped mangabeys in social contexts. They are based on movements of the mouth, eyebrows, and ears, possibly graded in intensity and produced independently or in combination. We showed that most of the facial displays were produced preferentially in specific social contexts and that repertoires varied with subjects' characteristics, highlighting the communicative function of these displays. Moreover, behavioral markers of intentionality commonly used in gestural studies were found to accompany the production of some of the facial signals observed. Particularly, playful "open mouth" appeared strongly associated with intentionality indices, as previously noticed in ape species. All other facial displays, except yawns, did not exhibit all defined intentional indices but were, at least, directed toward a recipient. Interestingly, yawns presented different variants of intensity associated presumably with different social functions. Altogether, these results emphasize the communicative function of red-capped mangabeys' facial displays and provide a basis for further research on their intentional communication. (PsycInfo Database Record (c) 2021 APA, all rights reserved).

Sensitivity to recipient's attention and responsiveness are critical markers of intentional communication. Although previous research showed that ape gestures can be intentional, few studies have yet addressed this question concerning monkeys. Here, we characterise the effect of a recipient's presence, attentional state and responsiveness on the interspecific gestural communication of captive red-capped mangabeys (Cercocebus torquatus). Previous reports showed that they produced learnt begging gestures towards a human recipient preferentially when the latter was facing them. We used here a novel setup that allows subjects to move around an experimenter and to use different modalities (visual and acoustic) to communicate. We found that when the recipient was not facing them, mangabeys moved to a position in the visual field of their recipient rather than using attention-getters. Interestingly, unlike apes, they did not elaborate their communication visually or acoustically when the experimenter did not respond favourably to their begging. However, our results may suggest that begging gestures were goal-directed, since mangabeys inhibited them when the experimenter was not available to answer immediately (i.e. give a reward). Overall, red-capped mangabeys' interspecific visual communication presented intentionality features, but their use of begging gestures was less flexible than that of great apes in similar situations.

Increased reproduction success, enhanced foraging and reduced predation risk are usually regarded as major factors favouring the evolution of social behaviour. Here we formulate a series of hypotheses relating sexual, ecological and behavioural factors to evaluate their explanatory value for 13 extant otter species, estimating the extent to which each factor contributes to the sociality of each species. We also compare individual behaviours within some of the species. Four otter species are obligatory social; four are obligatory solitary; five present both types of social organization. Social organizations of otter species are not related to their phylogenetic relationships. However, many otter species exhibit intra-species patterns of flexible social lifestyles. Both solitary and social otters adjust their social patterns in response chiefly to food availability, but also to habitat features and competition. Group living is more common when intraspecific competition is reduced or trophic resources replenish rapidly. Under these circumstances, group members often forage individually. When otters forage individually, they often switch prey type when they compete with other conspecifics. Social structures of otters fall into seven types: (1) family groups; (2) extended family groups, often with an alpha dominant pair; (3) highly social groups with helpers; (4) collective hunting groups; (5) solitary lifestyle; (6) unstable mixed-sex groups; and (7) single-sex bachelor groups. When an individual of a species with variable sociality adopts one type of sociality, this may be only temporary. Variations in social life are actually based on a series of events that induce individuals to make decisions taking ecological factors into account. Although ontogenetic factors can influence delayed dispersal of otters, social factors rather than ecological factors could play an important role in the formation of groups, and cohesiveness and kinship appear to be secondary effects of reduced dispersal more than primary causes for living in a group. Appropriate adjustment of group behaviour reduces the cost of sociality because individuals avoid social interactions when benefits are low but gather together when group living provides real advantages. Although any one model is unlikely to explicate all facets of sociality, evolution towards a social group results mainly from interactions within a family.

Studies on auditory laterality have revealed asymmetries for processing, particularly species-specific signals, in vertebrates and that each hemisphere may process different features according to their functional value. Processing of novel, intense emotion-inducing or finer individual features may require attention and we hypothesised that the functional pertinence of the stimuli may be modulating attentional processes and hence lateralisation of sound processing. Behavioural measures in (food) distracted captive Campbell's monkeys and electrophysiological recordings in anesthetised (versus awake) European starlings were performed during the broadcast of auditory stimuli with different functional saliences (e.g., familiar/novel). In Campbell's monkeys, only novel sounds elicited lateralised responses, with a right hemisphere preference. Unfamiliar sounds elicited more head movements, reflecting enhanced attention, whereas familiar (usual in the home environment) sounds elicited few responses, and thus might not be arousing enough to stimulate attention. In starlings, in field L, when awake, individual identity was processed more in the right hemisphere, whereas, when anaesthetised, the left hemisphere was more involved in processing potentially socially meaningless sounds. These results suggest that the attention-getting property of stimuli may be an adapted concept for explaining hemispheric auditory specialisation. An attention-based model may reconcile the different existing hypotheses of a Right Hemisphere-arousal/intensity or individual based lateralisation.

Comparative studies can help understand better brain functional lateralization for manipulation and language. This study investigated and compared, for the first time, human adults' laterality for manipulation and gestures in a non-experimental social context. We analysed the manual laterality of 48 beach volleyball athletes for four frequently expressed behaviours: a complex throwing action (jump serve) and three gestures (CLAP HAND, PUMP FIST and SLAP HAND-TO-HAND). We evaluated population-level laterality bias for each of the four behaviours separately, compared manual laterality between behaviours and investigated factors influencing gestural laterality. We furthered our between-gestures comparison by taking into account three categories of factors simultaneously: gesture characteristics (sensory modality), interactional context components (positions of interactants and emotional valence), and individual demographic characteristics (age, sex and country). Our study showed that (1) each behaviour considered presented a population-level right-hand bias, (2) differences of laterality between behaviours were probably related to gesture sensory modality and (3) signaller's laterality was modulated differently in relation to positions of interactants, emotional valence, age and sex. Our results support the literature suggesting that left-hemisphere specialization for manipulation and language (speech and gestures) may have evolved from complex manual activities such as throwing and from gestural communication.

Investigating in depth the mechanisms underlying human and non-human primate intentional communication systems (involving gestures, vocalisations, facial expressions and eye behaviours) can shed light on the evolutionary roots of language. Reports on non-human primates, particularly great apes, suggest that gestural communication would have been a crucial prerequisite for the emergence of language, mainly based on the evidence of large communication repertoires and their associated multifaceted nature of intentionality that are key properties of language. Such research fuels important debates on the origins of gestures and language. We review here three non-mutually exclusive processes that can explain mainly great apes' gestural acquisition and development: phylogenetic ritualisation, ontogenetic ritualisation, and learning via social negotiation. We hypothesise the following scenario for the evolutionary origins of gestures: gestures would have appeared gradually through evolution via signal ritualisation following the principle of derived activities, with the key involvement of emotional expression and processing. The increasing level of complexity of socioecological lifestyles and associated daily manipulative activities might then have enabled the acquisition and development of different interactional strategies throughout the life cycle. Many studies support a multimodal origin of language. However, we stress that the origins of language are not only multimodal, but more broadly multicausal. We propose a multicausal theory of language origins which better explains current findings. It postulates that primates' communicative signalling is a complex trait continually shaped by a cost-benefit trade-off of signal production and processing of interactants in relation to four closely interlinked categories of evolutionary and life cycle factors: species, individual and context-related characteristics as well as behaviour and its characteristics. We conclude by suggesting directions for future research to improve our understanding of the evolutionary roots of gestures and language.

Studying the relationships between the directions of brain lateralization for handedness and language can shed light on mechanisms underlying hemispheric specialization for manipulation and signalling functions. We investigated the influence of manipulation and communication functions and of recipient species (conspecific- versus human-directed communication) on manual laterality in signalling context, taking several factors into account simultaneously. We assessed laterality in 39 chimpanzees ( Pan troglodytes ), including 4 manipulators (mechanically effective social actions used to get things done) and 18 gestures (mechanically ineffective social actions implying that the signaller takes the recipient’s response into account). We focused on the following factors: interactional context components (e.g., visual fields of both interactants), degree of use of signals (“rare” for signals performed by only a few subjects in the population or “common” for signals performed by many subjects), mechanical effectiveness, subjects’ sociodemographic characteristics (e.g., age and hierarchy), and recipient species. We found a significant population-level right-hand bias for one type of human-directed gesture ( slap hand ). Mechanical effectiveness influenced laterality: right-hand use was more pronounced for conspecific-directed gestures than for conspecific-directed manipulators. The laterality of conspecific-directed gestures overall did not differ from that of human-directed gestures. However, we found an indirect influence of recipient species on laterality as conspecific- and human-directed gestural lateralities were modulated differently by the position of the recipient in the signaller’s visual field and by signaller’s age. We hypothesize that the communication nature of gestures might have developed from manipulators. Manipulators may have contributed to the emergence and the evolution of the left-lateralized communication system in primates.

Understanding variations of apes' laterality between activities is a central issue when investigating the evolutionary origins of human hemispheric specialization of manual functions and language. We assessed laterality of 39 chimpanzees in a non-communication action similar to termite fishing that we compared with data on five frequent conspecific-directed gestures involving a tool previously exploited in the same subjects. We evaluated, first, population-level manual laterality for tool-use in non-communication actions; second, the influence of sociodemographic factors (age, sex, group, and hierarchy) on manual laterality in both non-communication actions and gestures. No significant right-hand bias at the population level was found for non-communication tool use, contrary to our previous findings for gestures involving a tool. A multifactorial analysis revealed that hierarchy and age particularly modulated manual laterality. Dominants and immatures were more right-handed when using a tool in gestures than in non-communication actions. On the contrary, subordinates, adolescents, young and mature adults as well as males were more right-handed when using a tool in non-communication actions than in gestures. Our findings support the hypothesis that some primate species may have a specific left-hemisphere processing gestures distinct from the cerebral system processing non-communication manual actions and to partly support the tool use hypothesis.

This review highlights the scientific advances concerning the origins of human right‐handedness and language (speech and gestures). The comparative approach we adopted provides evidence that research on human and non‐human animals’ behavioural asymmetries helps understand the processes that lead to the strong human left‐hemisphere specialisation. We review four major non‐mutually exclusive environmental factors that are likely to have shaped the evolution of human and non‐human primates’ manual asymmetry: socioecological lifestyle, postural characteristics, task‐level complexity and tool use. We hypothesise the following scenario for the evolutionary origins of human right‐handedness: the right‐direction of modern humans’ manual laterality would have emerged from our ecological (terrestrial) and social (multilevel system) lifestyle; then, it would have been strengthened by the gradual adoption of the bipedal stance associated with bipedal locomotion, and the increasing level of complexity of our daily tasks including bimanual coordinated actions and tool use. Although hemispheric functional lateralisation has been shaped through evolution, reports indicate that many factors and their mutual intertwinement can modulate human and non‐human primates’ manual laterality throughout their life cycle: genetic and environmental factors, mainly individual sociodemographic characteristics (e.g., age, sex and rank), behavioural characteristics (e.g., gesture per se and gestural sensory modality) and context‐related characteristics (e.g., emotional context and position of target). These environmental (evolutionary and life cycle) factors could also have influenced primates’ manual asymmetry indirectly through epigenetic modifications. All these findings led us to propose the hypothesis of a multicausal origin of human right‐handedness.

A growing consensus favors the predominance of the human left hemisphere in manipulation and language (speech and gestures). However, the mechanisms underlying brain lateralization for noncommunication and communication functions are still unclear. Many studies emphasize the ambiguous nature of the relationship between the directions of brain lateralization for manipulation and for language. A comparative evolutionary approach investigating lateralization and communication mechanisms in our closest living relatives can improve our understanding of human hemispheric specialisation. We review theories concerning the evolutionary origins of human right-handedness and language and studies, highlighting the relevance of a comparative evolutionary approach. We discuss four methodological issues related to the study of handedness and communication: 1) categorisation of signaling based on criteria of intentionality; 2) use of a comprehensive multimodal and multifactorial approach; 3) investigation of laterality in intraspecific gestures; and 4) comparison of manual laterality between gestures and noncommunication actions. Deeper investigations of the multifaceted nature of intentionality in communication and the multidimensionality of manual laterality and communication in humans and other primates are needed. Finally, we make four recommendations to enhance our understanding of the evolutionary origins of human right-handedness and language: 1) implementing a comprehensive and integrated investigation of laterality and communication; 2) considering socioecologically relevant contexts; 3) investigating intraspecific aspects of communication in humans and other primates that vary in their degree of sociality; and 4) using appropriate, standardized methods of data collection and analysis.

Despite significant scientific advances, the nature of the left-hemispheric systems involved in language (speech and gesture) and manual actions is still unclear. To date, investigations of human laterality focused mainly on non-communication functions. Although gestural laterality data have been published for infants and children, relatively little is known about laterality of human gestural communication. This study investigated human laterality in depth considering non-communication manipulation actions and various gesture types involving hands, feet, face and ears. We constructed an online laterality questionnaire including 60 items related to daily activities. We collected 317 594 item responses by 5904 randomly selected participants. The highest percentages of strong left-lateralized (6.76%) and strong right-lateralized participants (75.19%) were for manipulation actions. The highest percentages of mixed left-lateralized (12.30%) and ambidextrous (50.23%) participants were found for head-related gestures. The highest percentage of mixed right-lateralized participants (55.33%) was found for auditory gestures. Every behavioural category showed a significant population-level right-side bias. More precisely, participants were predominantly right-lateralized for non-communication manual actions, for visual iconic, visual symbolic, visual deictic (with and without speech), tactile and auditory manual gestures as well as for podial and head-related gestures. Our findings support previous studies reporting that humans have left-brain predominance for gestures and complex motor activities such as tool-use. Our study shows that the Rennes Laterality Questionnaire is a useful research instrument to assess and analyse human laterality for both manipulation and communication functions.

A relevant approach to address the mechanisms underlying the emergence of the right-handedness/left-hemisphere language specialization of humans is to investigate both proximal and distal causes of language lateralization through the study of non-human primates' gestural laterality. We carried out the first systematic, quantitative comparison of within-subjects' and between-species' laterality by focusing on the laterality of intraspecific gestures of chimpanzees (Pan troglodytes) and gorillas (Gorilla gorilla) living in six different captive groups. We addressed the following two questions: (1) Do chimpanzees and gorillas exhibit stable direction of laterality when producing different types of gestures at the individual level? If yes, is it related to the strength of laterality? (2) Is there a species difference in gestural laterality at the population level? If yes, which factors could explain this difference? During 1356 observation hours, we recorded 42335 cases of dyadic gesture use in the six groups totalling 39 chimpanzees and 35 gorillas. Results showed that both species could exhibit either stability or flexibility in their direction of gestural laterality. These results suggest that both stability and flexibility may have differently modulated the strength of laterality depending on the species social structure and dynamics. Furthermore, a multifactorial analysis indicates that these particular social components may have specifically impacted gestural laterality through the influence of gesture sensory modality and the position of the recipient in the signaller's visual field during interaction. Our findings provide further support to the social theory of laterality origins proposing that social pressures may have shaped laterality through natural selection.

In contrast to human adults, risk proneness in the gain domain is usually observed in both young children and non-human primates. It is currently unclear what mechanism might be underlying such economic preferences. We investigated decision-making under risk of gain in toddlers and monkeys. The choices of 2.5-year-old children and red-capped mangabeys (Cercocebus torquatus torquatus) were examined in a gambling task for food reward in which participants have to choose between two options, a secure option and a risky option. In contrast to monkeys, toddlers showed a strong preference for the risky option over the safe option. In order to test the hypothesis that risky choices in participants reflect inhibitory control difficulties, toddlers and mangabeys were presented in Experiment 2 with a situation analogous to that used in Experiment 1 except for the fact that the opaque cover under which was placed the secure option was replaced by a transparent cover. In this second experiment, toddlers continued to show a preference for the risky option over the safe option. In contrast, mangabeys showed a preference for the safe option over the risky option in Trial 1 but they shifted their economic preferences in Trial 2. We argue that decision-making strategies under risk of gain in both toddlers and mangabeys (a) do not reflect poor behavioral control and (b) are not reducible to perception-action couplings.

Relationships between humans' manual laterality in non-communicative and communicative functions are still poorly understood. Recently, studies showed that chimpanzees' manual laterality is influenced by functional, interactional and individual factors and their mutual intertwinement. However, what about manual laterality in species living in stable social groups? We tackled this question by studying three groups of captive gorillas (N=35) and analysed their most frequent manual signals: three manipulators and 16 gesture types. Our multifactorial investigation showed that conspecific-directed gestures were overall more right-lateralized than conspecific-directed manipulators. Furthermore, it revealed a difference between conspecific- and human-directed gestural laterality for signallers living in one of the study groups. Our results support the hypothesis that gestural laterality is a relevant marker of language left-brain specialisation. We suggest that components of communication and of manipulation (not only of an object but also of a conspecific) do not share the same lateralised cerebral system in some primate species.

Many vertebrates present behavioural asymmetries (i.e. functional and/or structural specializations of left and right sides of the brain); however, evidence for arthropods is scarce. Some behavioural asymmetries can be correlated with morphology. A better understanding of behavioural asymmetries would be a crucial step to understand the evolution of brain asymmetries. Here, we investigated behavioural asymmetries of adult males of a mygalomorph Brachypelma albopilosum. First, as the time budgets of these active hunting spiders had not yet been documented, we established the temporal distribution of males' motor activity to be able to test them when they were active. Their motor activity peaks during the night and again early morning (around 3h and 10h local time). Then, choice tests in a T-maze assessed the effects of light and of odours separately. Our results revealed that male tarantulas' activity increased when they perceived light or the odour of conspecific females. Latencies to enter into the T-maze were shorter when a light cue was present and even shorter when odour cues (of prey or of conspecifics) were present. Choice between two identical cues (light or female odours) in a T-maze revealed, for the first time, their right behavioural laterality. Surprisingly, no significant external morphological differences could be evidenced between left and right eyes, lengths of the first legs or densities of mechanoreceptors and chemoreceptors on the tarsi of males' first legs to suggest perceptual asymmetry supporting this behavioural asymmetry. This is the first report concerning tarantulas' behavioural laterality.

Running comparative studies of laterality in mammals is a way to deepen our understanding of the evolution of the brain hemisphere functions. Studies on vision highlighted a possible task-sharing between hemispheres depending on the characteristics of the observers, the nature of the observed stimulus and the context of the observation, a phenomenon that could go beyond the monitoring of conspecifics. Cetaceans are predators that adapted to an aquatic habitat and display a clear crossing of fibers to the side of the brain opposite the eye of origin. Here, we analysed the interactions between humans and cetaceans when free-ranging orcas approach divers. Our study concentrated on the spontaneous exploratory behaviours of divers by orcas depending on their age and sex, and on the possible expression of a visual laterality. The results showed a significant preference for the use of the left eye but exclusively in adult females. Adult males had a more sustained attention than adult females, marked by a higher spatial proximity to divers, slower approaches and longer look durations. Adult females, probably more cautious, explored from the distance and more furtively. Our findings support a possible link between attentional/motivational states and visual laterality in mammals.