Search

Inhibitory control, the inhibition of impulsive behaviours, is believed to be key in navigating a complex social environment. Species characterised by higher social tolerance, living in more complex groups, with more diverse relationships, face higher uncertainty regarding the outcome of social interactions, and therefore, would benefit from employing more inhibitory strategies. To date, little is known about the selective forces that favour the evolution of inhibitory control. In this study, we compared inhibitory control skills in three closely related macaque species which differ in their social tolerance style. We tested 66 macaques from two institutions (Macaca mulatta, low tolerance; M. fascicularis, medium tolerance; and M. tonkeana, high tolerance) using a battery of validated inhibitory control touchscreen tasks. Higher social tolerance was associated with enhanced inhibitory control performances. More tolerant species were less impulsive and less distracted by pictures of unknown conspecifics. Interestingly, we did not find evidence that social tolerance degree was associated with performance in reversal learning. Overall, our results support the hypothesis that evolution has promoted the development of socio-cognitive skills to cope with the demands related to the complexity of the social environment.

A wealth of experimental and observational evidence suggests that faces have become increasingly important in the communication system of primates over evolutionary time and that both the static and moveable aspects of faces convey considerable information. Therefore, whenever there is a visual component to any multicomponent signal the face is potentially relevant. However, the role of the face is not always considered in primate multicomponent communication research. We review the literature and make a case for greater focus on the face going forward. We propose that the face can be overlooked for two main reasons: first, due to methodological difficulty. Examination of multicomponent signals in primates is difficult, so scientists tend to examine a limited number of signals in combination. Detailed examination of the subtle and dynamic components of facial signals is particularly hard to achieve in studies of primates. Second, due to a common assumption that the face contains “emotional” content. A priori categorisation of facial behavior as “emotional” ignores the potentially communicative and predictive information present in the face that might contribute to signals. In short, we argue that the face is central to multicomponent signals (and also many multimodal signals) and suggest future directions for investigating this phenomenon.

Ambiguity in communicative signals may lead to misunderstandings and thus reduce the effectiveness of communication, especially in unpredictable interactions such as between closely matched rivals or those with a weak social bond. Therefore, signals used in these circumstances should be less ambiguous, more stereotyped and more intense. To test this prediction, we measured facial movements of crested macaques ( Macaca nigra ) during spontaneous social interaction, using the Facial Action Coding System for macaques (MaqFACS). We used linear mixed models to assess whether facial movement intensity and variability varied according to the interaction outcome, the individuals' dominance relationship and their social bond. Movements were least intense and most variable in affiliative contexts, and more intense in interactions between individuals who were closely matched in terms of dominance rating. We found no effect of social bond strength. Our findings provide evidence for a reduction in ambiguity of facial behaviour in risky social situations but do not demonstrate any mitigating effect of social relationship quality. The results indicate that the ability to modify communicative signals may play an important role in navigating complex primate social interactions. This article is part of the theme issue ‘Cognition, communication and social bonds in primates’.

Darwin observed that form, and in his view, meaning, of facial behaviour (observable changes in the appearance of the face, often termed facial ‘expression’) is similar between a wide range of species and concluded that this must be due to a shared ancestral origin. Yet, as with all social behaviours, exactly how to define similarity and determine homology is debated. Facial behaviour is linked to specific facial muscle movements, so one important factor in determining homology is the anatomical basis of facial behaviours that appear similar in both appearance and social function. The Facial Action Coding System (FACS) was developed for the scientific measurement of human facial behaviour and is based on individual facial muscle movements (Ekman and Friesen, 1978). FACS has since been modified for use with various non-human primate species (chimpanzees, macaques, hylobatids, orangutans) and domestic species (dogs, cats, horses). These FACS can be used to trace continuity of form in facial behaviour across species and build a better understanding of the evolution of facial communication in mammals.

Human language is thought to have evolved from non-linguistic communication systems present in the primate lineage. Scientists rely on data from extant primate species to estimate how this happened, with debates centering around which modality (vocalization, gesture, facial expression) was a likely precursor. In 2011, we demonstrated that different theoretical and methodological approaches are used to collect data about each modality, rendering datasets incomplete and comparisons problematic. Here, 10 years later, we conducted a follow-up systematic review to test whether patterns have changed, examining the primate communication literature published between 2011 and 2020. In sum, despite the promising progress in addressing some gaps in our knowledge, systematic biases still exist and multimodal research remains uncommon. We argue that theories of language evolution are unlikely to advance until the field of primate communication research acknowledges and rectifies the gaps in our knowledge.

The NetFACS project is funded by a Leverhulme Trust Research Project Grant. AM received funding from the British Academy Newton International Fellowship scheme. Understanding facial signals in humans and other species is crucial for understanding the evolution, complexity, and function of the face as a communication tool. The Facial Action Coding System (FACS) enables researchers to measure facial movements accurately, but we currently lack tools to reliably analyse data and efficiently communicate results. Network analysis can provide a way to use the information encoded in FACS datasets: by treating individual AUs (the smallest units of facial movements) as nodes in a network and their co-occurrence as connections, we can analyse and visualise differences in the use of combinations of AUs in different conditions. Here, we present ‘NetFACS’, a statistical package that uses occurrence probabilities and resampling methods to answer questions about the use of AUs, AU combinations, and the facial communication system as a whole in humans and non-human animals. Using highly stereotyped facial signals as an example, we illustrate some of the current functionalities of NetFACS. We show that very few AUs are specific to certain stereotypical contexts; that AUs are not used independently from each other; that graph-level properties of stereotypical signals differ; and that clusters of AUs allow us to reconstruct facial signals, even when blind to the underlying conditions. The flexibility and widespread use of network analysis allows us to move away from studying facial signals as stereotyped expressions, and towards a dynamic and differentiated approach to facial communication. Publisher PDF

Objectives - While it has been demonstrated that even subtle variation in human facial expressions can lead to significant changes in the meaning and function of expressions, relatively few studies have examined primate facial expressions using similarly objective and rigorous analysis. Construction of primate facial expression repertoires may, therefore, be oversimplified, with expressions often arbitrarily pooled and/or split into subjective pigeonholes. Our objective is to assess whether subtle variation in primate facial expressions is linked to variation in function, and hence to inform future attempts to quantify complexity of facial communication.Materials and Methods - We used Macaque Facial Action Coding System, an anatomically based and hence more objective tool, to quantify “silent bared‐teeth” (SBT) expressions produced by wild crested macaques engaging in spontaneous behavior, and utilized discriminant analysis and bootstrapping analysis to look for morphological differences between SBT produced in four different contexts, defined by the outcome of interactions: Affiliation, Copulation, Play, and Submission.Results - We found that SBT produced in these contexts could be distinguished at significantly above‐chance rates, indicating that the expressions produced in these four contexts differ morphologically. We identified the specific facial movements that were typically used in each context, and found that the variability and intensity of facial movements also varied between contexts.Discussion - These results indicate that nonhuman primate facial expressions share the human characteristic of exhibiting meaningful subtle differences. Complexity of facial communication may not be accurately represented simply by building repertoires of distinct expressions, so further work should attempt to take this subtle variability into account.

The studies were funded by a Leverhulme Trust Research Project Grant Cultural variation in the social function and expression of guilt awarded to the seventh and fourth authors (RPG-2016-206) and the Leverhulme Trust Research Project Grant Rethinking complexity in facial communication systems awarded to the sixth author (RPG-2018-334). Guilt is a complex emotion with a potentially important social function of stimulating cooperative behaviours towards and from others, but whether the feeling of guilt is associated with a recognisable pattern of nonverbal behaviour is unknown. We examined the production and perception of guilt in two different studies, with a total of 238 participants with various places of origin. Guilt was induced experimentally, eliciting patterns of movement that were associated with both the participants’ self-reported feelings of guilt and judges’ impressions of their guilt. Guilt was most closely associated with frowning and neck touching. While there were differences between self-reported guilt and perception of guilt the findings suggest that there are consistent patterns that could be considered a non-verbal signal of guilt in humans. Publisher PDF

Self-directed behaviours in primates as a response to increasing psychological or physiological stress are a well-studied phenomenon. There is some evidence that these behaviours can be contagious when observed by conspecifics, but the adaptive function of this process is unclear. The ability to perceive stress in others and respond to it could be an important part of sustaining cohesiveness in social primates, but spontaneously acquiring stress-related behaviours (and potentially emotional states) from all group mates via contagion could be maladaptive. To investigate this, a group of captive Barbary macaques, Macaca sylvanus, were presented with videos of conspecifics engaging in self-directed behaviour (scratching) and neutral behaviour. Behavioural responses as a result of exposure to the stimuli were compared (1) between familiar and unfamiliar individuals, and (2) within familiar individuals to consider the modulating effects of social relationships. Our results did not show contagious scratching in this species. However, there were differences in how individuals attended to the scratching stimuli. Subjects were more attentive to scratching videos than to neutral videos and familiar than unfamiliar individuals. Within the familiar individuals, subjects were more attentive to those to whom they were weakly bonded. We suggest that increased attention to scratching behaviours may be adaptive in order to monitor and avoid stressed group mates, whose subsequent behaviour may be unpredictable and aggressive. Monitoring group mates who are not allies may also be adaptive as they may pose the biggest risk. These findings will help increase our understanding of subtle cues that can be communicative in primates, and also the evolutionary steps towards understanding others.

Language's intentional nature has been highlighted as a crucial feature distinguishing it from other communication systems. Specifically, language is often thought to depend on highly structured intentional action and mutual mindreading by a communicator and recipient. Whilst similar abilities in animals can shed light on the evolution of intentionality, they remain challenging to detect unambiguously. We revisit animal intentional communication and suggest that progress in identifying analogous capacities has been complicated by (i) the assumption that intentional (that is, voluntary) production of communicative acts requires mental-state attribution, and (ii) variation in approaches investigating communication across sensory modalities. To move forward, we argue that a framework fusing research across modalities and species is required. We structure intentional communication into a series of requirements, each of which can be operationalised, investigated empirically, and must be met for purposive, intentionally communicative acts to be demonstrated. Our unified approach helps elucidate the distribution of animal intentional communication and subsequently serves to clarify what is meant by attributions of intentional communication in animals and humans.

Facial displays (or expressions) are a primary means of visual communication among conspecifics in many mammalian orders. Macaques are an ideal model among primates for investigating the co-evolution of facial musculature, facial displays, and social group size/behavior under the umbrella of "ecomorphology". While all macaque species share some social behaviors, dietary, and ecological parameters, they display a range of social dominance styles from despotic to tolerant. A previous study found a larger repertoire of facial displays in tolerant macaque species relative to despotic species. The present study was designed to further explore this finding by comparing the gross morphological features of mimetic muscles between the Sulawesi macaque (Macaca nigra), a tolerant species, and the rhesus macaque (M. mulatta), a despotic species. Five adult M. nigra heads were dissected and mimetic musculature was compared to those from M. mulatta. Results showed that there was general similarity in muscle presence/absence between the species as well as muscle form except for musculature around the external ear. M. mulatta had more musculature around the external ear than M. nigra. In addition, M. nigra lacked a zygomaticus minor while M. mulatta is reported to have one. These morphological differences match behavioral observations documenting a limited range of ear movements used by M. nigra during facial displays. Future studies focusing on a wider phylogenetic range of macaques with varying dominance styles may further elucidate the roles of phylogeny, ecology, and social variables in the evolution of mimetic muscles within Macaca Anat Rec, 299:1317-1324, 2016. © 2016 Wiley Periodicals, Inc.

There is widespread acceptance that facial expressions are useful in social interactions, but empirical demonstration of their adaptive function has remained elusive. Here, we investigated whether macaques can use the facial expressions of others to predict the future outcomes of social interaction. Crested macaques (Macacanigra) were shown an approach between two unknown individuals on a touchscreen and were required to choose between one of two potential social outcomes. The facial expressions of the actors were manipulated in the last frame of the video. One subject reached the experimental stage and accurately predicted different social outcomes depending on which facial expressions the actors displayed. The bared-teeth display (homologue of the human smile) was most strongly associated with predicted friendly outcomes. Contrary to our predictions, screams and threat faces were not associated more with conflict outcomes. Overall, therefore, the presence of any facial expression (compared to neutral) caused the subject to choose friendly outcomes more than negative outcomes. Facial expression in general, therefore, indicated a reduced likelihood of social conflict. The findings dispute traditional theories that view expressions only as indicators of present emotion and instead suggest that expressions form part of complex social interactions where individuals think beyond the present. Electronic supplementary material The online version of this article (doi:10.1007/s10071-016-0992-3) contains supplementary material, which is available to authorized users.

Mammals use their faces in social interactions more so than any other vertebrates. Primates are an extreme among most mammals in their complex, direct, lifelong social interactions and their frequent use of facial displays is a means of proximate visual communication with conspecifics. The available repertoire of facial displays is primarily controlled by mimetic musculature, the muscles that move the face. The form of these muscles is, in turn, limited by and influenced by phylogenetic inertia but here we use examples, both morphological and physiological, to illustrate the influence that social variables may exert on the evolution and form of mimetic musculature among primates. Ecomorphology is concerned with the adaptive responses of morphology to various ecological variables such as diet, foliage density, predation pressures, and time of day activity. We present evidence that social variables also exert selective pressures on morphology, specifically using mimetic muscles among primates as an example. Social variables include group size, dominance ‘style’, and mating systems. We present two case studies to illustrate the potential influence of social behavior on adaptive morphology of mimetic musculature in primates: (1) gross morphology of the mimetic muscles around the external ear in closely related species of macaque (Macaca mulatta and Macaca nigra) characterized by varying dominance styles and (2) comparative physiology of the orbicularis oris muscle among select ape species. This muscle is used in both facial displays/expressions and in vocalizations/human speech. We present qualitative observations of myosin fiber‐type distribution in this muscle of siamang (Symphalangus syndactylus), chimpanzee (Pan troglodytes), and human to demonstrate the potential influence of visual and auditory communication on muscle physiology. In sum, ecomorphologists should be aware of social selective pressures as well as ecological ones, and that observed morphology might reflect a compromise between the demands of the physical and the social environments.

Primate facial expression has long been studied within a framework of emotion that has heavily influenced both theoretical approaches and scientific methods. For example, our understanding of the adaptive function and cognition of facial expression is tied to the assumption that facial expression is accompanied by an emotional internal state, which is decipherable by others. Here, we challenge this view and instead support the alternative that facial expression should also be conceptualised as an indicator of future behaviour as opposed to current emotional state alone (Behavioural Ecology View, Fridlund, 1994). We also advocate the use of standardised, objective methodology Facial Action Coding System, to avoid making assumptions about the underlying emotional state of animals producing facial expressions. We argue that broadening our approach to facial expression in this way will open new avenues to explore the underlying neurobiology, cognition and evolution of facial communication in both human and non-human primates.

Primates (including humans) scratch when stressed. So far, such scratching has been seen as a by-product of physiological processes associated with stress, and attributed proximate, regulatory function. However, it is possible that others could use this relationship between scratching and stress as an indication of the animal’s stress state, and thus scratching could potentially have social function. As a test of this theory, we measured the production of, and social responses to scratching in a group of free-ranging rhesus macaques (Macaca mulatta). Firstly, we found that the likelihood of scratching was greater around periods of heightened social stress, such as being in proximity to high-ranking individuals, or non-friends. Secondly, when macaques scratched, subsequent interactions were less likely to be aggressive and more likely to be affiliative. Potential attackers may avoid attacking stressed individuals as stressed individuals could behave unpredictably or be weakened by their state of stress (rendering aggression risky and/or unnecessary). Observable stress behaviour could therefore have additional adaptive value by reducing the potential for escalated aggression, benefiting both senders and receivers by facilitating social cohesion. This basic ability to recognise stress in others could also be an important component in the evolution of social cognition such as empathy.

The domestic cat (Felis silvestris catus) is quickly becoming the most popular animal companion in the world. The evolutionary processes that occur during domestication are known to have wide effects on the morphology, behaviour, cognition and communicative abilities of a species. Since facial expression is central to human communication, it is possible that cat facial expression has been subject to selection during domestication. Standardised measurement techniques to study cat facial expression are, however, currently lacking. Here, as a first step to enable cat facial expression to be studied in an anatomically based and objective way, CatFACS (Cat Facial Action Coding System) was developed. Fifteen individual facial movements (Action Units), six miscellaneous movements (Action Descriptors) and seven Ear Action Descriptors were identified in the domestic cat. CatFACS was then applied to investigate the impact of cat facial expression on human preferences in an adoption shelter setting. Rehoming speed from cat shelters was used as a proxy for human selective pressure. The behaviour of 106 cats ready for adoption in three different shelters was recorded during a standardised encounter with an experimenter. This experimental setup aimed to mimic the first encounter of a cat with a potential adopter, i.e. an unfamiliar human. Each video was coded for proximity to the experimenter, body movements, tail movements and face movements. Cat facial movements were not related to rehoming speed, suggesting that cat facial expression may not have undergone significant selection. In contrast, rubbing frequency was positively related to rehoming speed. The findings suggest that humans are more influenced by overt prosocial behaviours than subtle facial expression in domestic cats.

Many facilities that house captive primates play music for animal enrichment or for caregiver enjoyment. However, the impact on primates is unknown as previous studies have been inconclusive. We conducted three studies with zoo-housed chimpanzees (Pan troglodytes) and one with group-housed chimpanzees at the National Centre for Chimpanzee Care to investigate the effects of classical and pop/rock music on various variables that may be indicative of increased welfare. Study one compared the behaviour and use of space of 18 animals when silence, classical or pop/rock music was played into one of several indoor areas. Overall, chimpanzees did not actively avoid the area when music was playing but were more likely to exit the area when songs with higher beats per minute were broadcast. Chimpanzees showed significantly fewer active social behaviours when music, rather than silence, was playing. They also tended to be more active and engage in less abnormal behaviour during the music but there was no change to either self-grooming or aggression between music and silent conditions. The genre of music had no differential effects on the chimpanzees' use of space and behaviour. In the second study, continuous focal observations were carried out on three individuals with relatively high levels of abnormal behaviour. No differences in behaviour between music and silence periods were found in any of the individuals. The final two studies used devices that allowed chimpanzees to choose if they wanted to listen to music of various types or silence. Both studies showed that there were no persistent preferences for any type of music or silence. When taken together, our results do not suggest music is enriching for group-housed captive chimpanzees, but they also do not suggest that music has a negative effect on welfare.

Animals communicate using a vast array of different signals in different modalities. For chimpanzees, vocalizations, gestures and facial expressions are all important forms of communication, yet these signals have rarely been studied together holistically. The current study aimed to provide the first comprehensive repertoire of flexibly combined (‘free’) multimodal (MM) signals, and assess individual and contextual factors influencing production of, and responses to, unimodal (UM) and MM signals in wild chimpanzees. In total, 48 different free MM signals were produced. MM signals were produced at a significantly lower rate than UM signals, but 22 of 26 focal animals were observed to produce free MM signals. The relative production rates of different types of UM and MM signals differed significantly between the behavioural contexts investigated, showing flexible use of signals across contexts. In contrast, individual factors such as age, sex or rank of signaller did not appear to influence the type of signal produced or the likelihood of eliciting a response. Finally, we compared recipient responses to free MM grunt-gesture signals and matched UM component signals and found that these MM signals were more likely to elicit a response than a grunt alone, but were as likely to elicit a response as the gesture alone. The overall findings point to a widespread capacity for wild chimpanzees to flexibly combine signals from different modalities and highlight the importance of adopting a multimodal approach to studying communication.

Little is known about facial communication of lesser apes (family Hylobatidae) and how their facial expressions (and use of) relate to social organization. We investigated facial expressions (defined as combinations of facial movements) in social interactions of mated pairs in five different hylobatid species belonging to three different genera using a recently developed objective coding system, the Facial Action Coding System for hylobatid species (GibbonFACS). We described three important properties of their facial expressions and compared them between genera. First, we compared the rate of facial expressions, which was defined as the number of facial expressions per units of time. Second, we compared their repertoire size, defined as the number of different types of facial expressions used, independent of their frequency. Third, we compared the diversity of expression, defined as the repertoire weighted by the rate of use for each type of facial expression. We observed a higher rate and diversity of facial expression, but no larger repertoire, in Symphalangus (siamangs) compared to Hylobates and Nomascus species. In line with previous research, these results suggest siamangs differ from other hylobatids in certain aspects of their social behavior. To investigate whether differences in facial expressions are linked to hylobatid socio‐ecology, we used a Phylogenetic General Least Square (PGLS) regression analysis to correlate those properties with two social factors: group‐size and level of monogamy. No relationship between the properties of facial expressions and these socio‐ecological factors was found. One explanation could be that facial expressions in hylobatid species are subject to phylogenetic inertia and do not differ sufficiently between species to reveal correlations with factors such as group size and monogamy level.

Pseudoreplication (the pooling fallacy) is a widely acknowledged statistical error in the behavioural sciences. Taking a large number of data points from a small number of animals creates a false impression of a better representation of the population. Studies of communication may be particularly prone to artificially inflating the data set in this way, as the unit of interest (the facial expression, the call or the gesture) is a tempting unit of analysis. Primate communication studies (551) published in scientific journals from 1960 to 2008 were examined for the simplest form of pseudoreplication (taking more than one data point from each individual). Of the studies that used inferential statistics, 38% presented at least one case of pseudoreplicated data. An additional 16% did not provide enough information to rule out pseudoreplication. Generalized linear mixed models determined that one variable significantly increased the likelihood of pseudoreplication: using observational methods. Actual sample size (number of animals) and year of publication were not associated with pseudoreplication. The high prevalence of pseudoreplication in the primate communication research articles, and the fact that there has been no decline since key papers warned against pseudoreplication, demonstrates that the problem needs to be more actively addressed.

Many nonhuman animals produce facial expressions which sometimes bear clear resemblance to the facial expressions seen in humans. An understanding of this evolutionary continuity between species, and how this relates to social and ecological variables, can help elucidate the meaning, function, and evolution of facial expression. This aim, however, requires researchers to overcome the theoretical and methodological differences in how human and nonhuman facial expressions are approached. Here, we review the literature relating to nonhuman facial expressions and suggest future directions that could facilitate a better understanding of facial expression within an evolutionary context.

Comparing homologous expressions between species can shed light on the phylogenetic and functional changes that have taken place during evolution. To assess homology across species we must approach primate facial expressions in an anatomical, systematic, and standardized way. The Facial Action Coding System (FACS), a widely used muscle-based tool for analyzing human facial expressions, has recently been adapted for chimpanzees (Pan troglodytes: ChimpFACS), rhesus macaques (Macaca mulatta: MaqFACS), and gibbons (GibbonFACS). Here, we present OrangFACS, a FACS adapted for orangutans (Pongo spp.). Orangutans are the most arboreal and the least social great ape, so their visual communication has been assumed to be less important than vocal communication and is little studied. We scrutinized the facial anatomy of orangutans and coded videos of spontaneous orangutan behavior to identify independent movements: Action Units (AUs) and Action Descriptors (ADs). We then compared these facial movements with movements of homologous muscles in humans, chimpanzees, macaques, and gibbons. We also noted differences related to sexual dimorphism and developmental stages in orangutan facial morphology. Our results show 17 AUs and 7 ADs in orangutans, indicating an overall facial mobility similar to that found in chimpanzees, macaques, and gibbons but smaller than that found in humans. This facial movement capacity in orangutans may be the result of several, nonmutually exclusive explanations, including the need for facial communication in specialized contexts, phylogenetic inertia, and allometric effects.

Primates’ communicative signals are often dynamic and composed of multiple components, sometimes belonging to different sensory modalities. Such multicomponent signals are of crucial importance in the study of communication: the addition of extra features to a signal has the potential to modulate or change the meaning and message of the specific signal. Traditionally, however, components of compound signals are studied in isolation from each other, or communicative displays are studied as static and invariant wholes. Both approaches may not allow us to assess the full function of the signals. In crested macaques (Macaca nigra), the lipsmack (a display mainly used in affiliative interactions) can be produced alone or combined with other visual and acoustic communicative features. We investigated whether the composition of the lipsmack influenced the outcome of social interaction while controlling for relationship quality. Our results show that lipsmacks composed of both visual and vocal components (i.e. multimodal signals) increased the probability of affiliative contact, and therefore have enhanced signal value. The total number of visual components involved in the display had no effect but some visual components seemed more influential than others. By analyzing lipsmacking behavior as a composite, dynamic display, we were able to reveal a level of complexity that is not apparent when looking at each component separately. The results highlight the importance of a more integrative, multimodal approach to the study of primate communication.

We examined the effect of asking unanticipated interview questions on lie detection. We set up a room in which truth tellers (N = 40) performed five tasks. Liars (N = 40) observed the activity room through a window and were asked to generate a story in which they claim to have performed five tasks. The interview schedule included four thematically related pairs of questions. The first question of each pairing was easy for interviewees to anticipate, as it reflected how events are typically recalled. However, the second included a perspective shift (temporal or spatial), which made the question harder to anticipate. We also measured cognitive load via performance on an object sorting secondary task that interviewees undertook during interview. Compared with truth tellers, liars' verbal responses showed a greater decline in details across three out of four question pairs. Liars also sorted significantly fewer objects per minute on the secondary task.

The evolution and function of human and nonhuman primate facial expression can be better understood through multispecies comparative analyses. Anatomically based coding systems (Facial Action Coding Systems [FACS]) are developed to enable such analyses because they are standardized and systematic and aid identification of homologous expressions underpinned by similar muscle contractions. To date, FACS has been developed for humans, and subsequently modified for chimpanzees and rhesus macaques. Through detailed examination of facial anatomy and spontaneous movement, here we present a FACS for hylobatids (gibbons and siamangs): apes living in small groups and often thought not to display extensive facial movement. We identified and described 18 independent facial movements (Action Units and Action Descriptors) in the hylobatid face, and compared to humans, chimpanzees, and rhesus macaques. The findings show that the facial movement capacity of hylobatids bears great similarity to species living in much larger groups. The findings could be explained by phylogenetic inertia (especially because pair-bonding is likely to have emerged from social living) and/or the social demands of pair-bonding.

Gaze following, the ability to follow the direction in which others are looking, is thought to allow individuals to acquire valuable information from their physical and social environment. Recent studies, using artificial stimuli, showed that gaze following can be modulated by social factors such as dominance or social context, suggesting the importance of integrating these factors in future cognitive studies to understand better the evolution and function of gaze following. Whether this finding still holds true when animals are tested with conspecifics is unknown. Moreover, additional social factors remain to be tested. Testing live conspecifics, we show that friendship (i.e. strong positive bonds between individuals) improved gaze-following responses in a tolerant primate: the crested macaque. Subjects did not follow the gaze of friends more often, but in dyads characterized by a high friendship index, subjects were quicker to react to gaze cues. The increased social tolerance characteristic of crested macaques’ social relationships may lessen the constraints imposed by dominance and kinship, thus allowing sociocognitive abilities to be better used among friends. Together with previous findings, our results suggest the importance of relationship quality and species’ social style in shaping primate cognition.

Primate facial expressions (FEs) likely play an important role in primate society: through facial signals, individuals can potentially send and receive information and may benefit from coordinating their behavior accordingly. Many primates use a relaxed open mouth (ROM) facial display or “play face” (PF) during play behavior, where the mouth is open but teeth are covered. In addition to this conventional PF, however, Western Lowland gorillas (Gorilla gorilla gorilla) also use a full PF where the upper teeth are exposed. As the teeth are similarly exposed in the bared-teeth expression (which is a signal of appeasement, submission and/or affiliation), the full PF may be a blend of the PF and bared-teeth face, and have a different signal function to the PF alone. Focal animal sampling of captive Western Lowland gorillas (N = 10) showed that the full PF was more often observed in intense rather than gentle play, and intense play bouts that featured the full PF were longer than those that featured only the PF. Both expressions were associated with an increase in affinitive behavior between sender and receiver postplay, but only the full PF was associated with an increase higher than that of play alone. Overall, the findings suggest that the full PF has an additional role in coordinating and maintaining play, possibly though reducing uncertainty in the receiver and confirming that play is only play.

Individuals in large groups do not always contribute equally to group decision making. This may be due to constraints on conversational group size, as when a group is comprised of more than 4 people, it spontaneously fissions into smaller groups within which conversations take place. Thus, if conversations are attempted in larger groups, some individuals will not be part of the spontaneously forming discursive subgroup and, consequently, may not contribute to group decision making. Here, using a mock jury paradigm, the effect of hierarchically subdividing groups to mimic the spontaneous formation of optimally sized conversational groups (while maintaining an overarching group size of 12) was tested. Individuals in the subdivided condition reported greater equality of contribution to the decision-making process than individuals in the control condition, and experienced less inhibition from participation due to the activities of others.

For far too long, the behaviour of our pets has been seen (at best) as unconstrained by comprehensible evolutionary processes, or (at worst) entirely constructed by anthropomorphism. John Bradshaw’s Cat Sense is a welcome attempt to challenge this status quo by demonstrating that science can help us understand why domestic cats do what they do, and how they inveigled their way into our homes. Compared to their rival, the dog, however, we know very little about our cat guests, a fact that permeates through this book loud and clear.

Over 125 years ago, Charles Darwin (1872) suggested that the only way to fully understand the form and function of human facial expression was to make comparisons with other species. Nevertheless, it has been only recently that facial expressions in humans and related primate species have been compared using systematic, anatomically based techniques. Through this approach, large-scale evolutionary and phylogenetic analyses of facial expressions, including their homology, can now be addressed. Here, the development of a muscular-based system for measuring facial movement in rhesus macaques (Macaca mulatta) is described based on the well-known FACS (Facial Action Coding System) and ChimpFACS. These systems describe facial movement according to the action of the underlying facial musculature, which is highly conserved across primates. The coding systems are standardized; thus, their use is comparable across laboratories and study populations. In the development of MaqFACS, several species differences in the facial movement repertoire of rhesus macaques were observed in comparison with chimpanzees and humans, particularly with regard to brow movements, puckering of the lips, and ear movements. These differences do not seem to be the result of constraints imposed by morphological differences in the facial structure of these three species. It is more likely that they reflect unique specializations in the communicative repertoire of each species.

Insights into the character of social relations are rare but exciting glimpses when it comes to archaeological populations. This paper uses a multidisciplinary combination of archaeological and evolutionary biological approaches to assess the social value of the bared‐teeth motif (BTM) in pre‐Columbian iconography of the Greater Antilles. Bared teeth feature on discrete artifact categories within the material culture of many different groups in the Taino culture area after AD 1000. This BTM, usually part of facial anthropomorphic or zoomorphic decoration, is used for bodily adornments and items associated with healing and shamanic practices. It has generally been interpreted as representative of death, aggression, or the shamanic trance. However, the motif has never been examined in terms of its signal value as a positive facial expression. Studies of facial expression in human and nonhuman primates have shown that the bared‐teeth expression is used in social contexts as an unambiguous signal of nonaggres...

Research considering the effect of Presidential speech making style shows individuals are likely to attend to and be emotionally affected by the facial expressions of leaders, especially those that are inappropriate for the context. In this study we tested how rapid and subtle expressions of facial affect (i.e., expressions of less than one-second, often termed micro-expressions) in speeches by a political leader impacts participants’ emotional state. We do this by removing seven brief expressions (less than one-second) from a nearly twelve minute televised speech by President George H. W. Bush concerning the 1990 commitment of US military in response to Iraq’s invasion of Kuwait. FACS coding identified all these expressions as containing a component of smiling (lip corner puller: AU12), in some cases with the addition of other facial movements (sometimes associated with anger and/or disgust). Experiments carried out on 206 university undergraduates showed emotional state was altered as a result of these micro-expressions when the control (unaltered speech) and experimental group (micro-expressions removed from speech) were compared. Specifically, participants who viewed the micro-expressions felt less threatened and less angry. Thus, facial expression (even very brief micro-expressions) can have a significant impact on the receiver of a political speech.

The ability of odors to cue vivid and emotionally intense memories is well-known. However, the majority of research has focused on the extent to which odors can act as environmental cues to memory, where odors are presented alongside the stimuli to be remembered, rather than the extent to which pre-existing associations between odor and odor-related stimuli might influence memory. In this study, participants (n = 45 females in each experiment) were presented with words (two groups of odor-associated words and one neutral) on a computer screen and randomly assigned to one of three conditions where they recalled the words while inhaling from a bottle either rosemary, jasmine, or no odor (experiment 1) and peppermint, bergamot, or no odor (experiment 2). In experiment 2, participants then completed a lexical decision task (LDT). Experiment 1 revealed that, for those in the rosemary group, significantly more rosemary versus jasmine and neutral words were recalled. Experiment 2 replicated this effect for peppermint, though no odor-congruent effects were found in the LDT. These findings demonstrate that certain odors are able to cue memory for odor-associated words. Results are discussed in relation to connected odor association research and possible theoretical frameworks to account for these findings.