Slow development of vocal sequences through ontogeny in wild chimpanzees (Pan troglodytes verus)

The development of the unique, hierarchical, and endless combinatorial capacity in a human language requires neural maturation and learning through childhood. Compared with most non-human primates, where combinatorial capacity seems limited, chimpanzees present a complex vocal system comprising hundreds of vocal sequences. We investigated how such a complex vocal system develops and the processes involved. We recorded 10,929 vocal utterances of 98 wild chimpanzees aged 0–55 years, from Taï National Park, Ivory Coast. We developed customized Generalized non-Linear Models to estimate the ontogenetic trajectory of four structural components of vocal complexity: utterance length, diversity, probability of panting (requiring phonation across inhalation and exhalation), and probability of producing two adjacent panted units. We found chimpanzees need 10 years to reach adult levels of vocal complexity. In three variables, the steepest increase coincided with the age of first non-kin social interactions (2–5 years), and plateaued in sub-adults (8–10 years), as individuals integrate into adult social life. Producing two adjacent panted units may require more neuromuscular coordination of the articulators, as its emergence and steepest increase appear later in development. These results suggest prolonged maturational processes beyond those hitherto thought likely in species that do not learn their vocal repertoire. Our results suggest that multifaceted ontogenetic processes drive increases in vocal structural complexity in chimpanzees, particularly increases in social complexity and neuro-muscular maturation. As humans live in a complex social world, empirical support for the “social complexity hypothesis” may have relevance for theories of language evolution.

Research Highlights

• Chimpanzees need around 10 years to develop the vocal structural complexity present in the adult repertoire, way beyond the age of emergence of every single vocal unit.
• Multifaceted ontogenetic processes may drive increases in vocal structural complexity in chimpanzees, particularly increases in social complexity and neuro-muscular maturation.
• Non-linear increases in vocal complexity coincide with social developmental milestones.
• Vocal sequences requiring rapid articulatory change emerge later than other vocal sequences, suggesting neuro-muscular maturational processes continue through the juvenile years.

     

Quantifying latent social motivation and its associations with joint attention and language in infants at high and low likelihood for autism spectrum disorder

Social motivation—the psychobiological predisposition for social orienting, seeking social contact, and maintaining social interaction—manifests in early infancy and is hypothesized to be foundational for social communication development in typical and atypical populations. However, the lack of infant social-motivation measures has hindered delineation of associations between infant social motivation, other early-arising social abilities such as joint attention, and language outcomes. To investigate how infant social motivation contributes to joint attention and language, this study utilizes a mixed longitudinal sample of 741 infants at high (HL = 515) and low (LL = 226) likelihood for ASD. Using moderated nonlinear factor analysis (MNLFA), we incorporated items from parent-report measures to establish a novel latent factor model of infant social motivation that exhibits measurement invariance by age, sex, and familial ASD likelihood. We then examined developmental associations between 6- and 12-month social motivation, joint attention at 12–15 months, and language at 24 months of age. On average, greater social-motivation growth from 6–12 months was associated with greater initiating joint attention (IJA) and trend-level increases in sophistication of responding to joint attention (RJA). IJA and RJA were both positively associated with 24-month language abilities. There were no additional associations between social motivation and future language in our path model. These findings substantiate a novel, theoretically driven approach to modeling social motivation and suggest a developmental cascade through which social motivation impacts other foundational skills. These findings have implications for the timing and nature of intervention targets to support social communication development in infancy.

Highlights

• We describe a novel, theoretically based model of infant social motivation wherein multiple parent-reported indicators contribute to a unitary latent social-motivation factor.
• Analyses revealed social-motivation factor scores exhibited measurement invariance for a longitudinal sample of infants at high and low familial ASD likelihood.
• Social-motivation growth from ages 6–12 months is associated with better 12−15-month joint attention abilities, which in turn are associated with greater 24-month language skills.
• Findings inform timing and targets of potential interventions to support healthy social communication in the first year of life.

     

Auditory–visual intermodal matching based on individual recognition in a chimpanzee (Pan troglodytes)

The ability to recognize familiar individuals with different sensory modalities plays an important role in animals living in complex physical and social environments. Individual recognition of familiar individuals was studied in a female chimpanzee named Pan. In previous studies, Pan learned an auditory–visual intermodal matching task (AVIM) consisting of matching vocal samples with the facial pictures of corresponding vocalizers (humans and chimpanzees). The goal of this study was to test whether Pan was able to generalize her AVIM ability to new sets of voice and face stimuli, including those of three infant chimpanzees. Experiment 1 showed that Pan performed intermodal individual recognition of familiar adult chimpanzees and humans very well. However, individual recognition of infant chimpanzees was poorer relative to recognition of adults. A transfer test with new auditory samples (Experiment 2) confirmed the difficulty in recognizing infants. A remaining question was what kind of cues were crucial for the intermodal matching. We tested the effect of visual cues (Experiment 3) by introducing new photographs representing the same chimpanzees in different visual perspectives. Results showed that only the back view was difficult to recognize, suggesting that facial cues can be critical. We also tested the effect of auditory cues (Experiment 4) by shortening the length of auditory stimuli, and results showed that 200 ms vocal segments were the limit for correct recognition. Together, these data demonstrate that auditory–visual intermodal recognition in chimpanzees might be constrained by the degree of exposure to different modalities and limited to specific visual cues and thresholds of auditory cues.     

Beneficial effects of a music listening intervention on neural speech processing in 0–28-month-old children at risk for dyslexia

Familial risk for developmental dyslexia can compromise auditory and speech processing and subsequent language and literacy development. According to the phonological deficit theory, supporting phonological development during the sensitive infancy period could prevent or ameliorate future dyslexic symptoms. Music is an established method for supporting auditory and speech processing and even language and literacy, but no previous studies have investigated its benefits for infants at risk for developmental language and reading disorders. We pseudo-randomized N∼150 infants at risk for dyslexia to vocal or instrumental music listening interventions at 0–6 months, or to a no-intervention control group. Music listening was used as an easy-to-administer, cost-effective intervention in early infancy. Mismatch responses (MMRs) elicited by speech-sound changes were recorded with electroencephalogram (EEG) before (at birth) and after (at 6 months) the intervention and at a 28 months follow-up. We expected particularly the vocal intervention to promote phonological development, evidenced by enhanced speech-sound MMRs and their fast maturation. We found enhanced positive MMR amplitudes in the vocal music listening intervention group after but not prior to the intervention. Other music activities reported by parents did not differ between the three groups, indicating that the group effects were attributable to the intervention. The results speak for the use of vocal music in early infancy to support speech processing and subsequent language development in infants at developmental risk.

Research Highlights

• Dyslexia-risk infants were pseudo-randomly assigned to a vocal or instrumental music listening intervention at home from birth to 6 months of age.
• Neural mismatch responses (MMRs) to speech-sound changes were enhanced in the vocal music intervention group after but not prior to the intervention.
• Even passive vocal music listening in early infancy can support phonological development known to be deficient in dyslexia-risk.

     

Who's my little monkey? Effects of infant‐directed speech on visual retention in infant rhesus macaques

Both human and nonhuman primate adults use infant‐directed facial and vocal expressions across many contexts when interacting with infants (e.g., feeding, playing). This infant‐oriented style of communication, known as infant‐directed speech (IDS), seems to benefit human infants in numerous ways, including facilitating language acquisition. Given the variety of contexts in which adults use IDS, we hypothesized that IDS supports learning beyond the linguistic domain and that these benefits may extend to nonhuman primates. We exposed 2.5‐month‐old rhesus macaque infants (N = 15) to IDS, adult‐directed speech (ADS), and a non‐social control (CTR) during a video presentation of unrelated stimuli. After a 5‐ or 60‐minute delay, infants were shown the familiar video side‐by‐side with a novel video. Infants exhibited a novelty preference after the 5‐minute delay, but not after the 60‐minute delay, in the ADS and CTR conditions, and a novelty preference in the IDS condition only after the 60‐minute delay. These results are the first to suggest that exposure to IDS affects infants’ long‐term memory, even in non‐linguistic animals.     

Early social deprivation negatively affects social skill acquisition in chimpanzees (Pan troglodytes)

In a highly social species like chimpanzees, the process by which individuals become attuned to their social environment may be of vital importance to their chances of survival. Typically, this socialization process, defined by all acquisition experiences and fine-tuning efforts of social interaction patterns during ontogeny, occurs in large part through parental investment. In this study, we investigated whether maternal presence enhances the socialization process in chimpanzees by comparing the social interactions of orphaned and mother-reared individuals at the Chimfunshi Wildlife Orphanage Trust in Zambia. As response variables, we selected social interactions during which an elaborate level of fine-tuning is assumed to be necessary for sustaining the interaction and preventing escalation: social play. Comparing orphaned (n = 8) to sex- and age-matched mother-reared juvenile chimpanzees (n = 9), we hypothesized that the orphaned juveniles would play less frequently than the mother-reared and would be less equipped for fine-tuning social play (which we assayed by rates of aggression) because of the lack of a safe and facilitating social environment provided by the mother. First, contrary to our hypothesis, results showed that the orphaned juveniles engaged in social play more frequently than the mother-reared juveniles, although for significantly shorter amounts of time. Second, in support of our hypothesis, results showed that social play of the orphaned juveniles more often resulted in aggression than social play of the mother-reared juveniles. In conjunction, these results may indicate that, just like in humans, chimpanzee mothers provide their offspring with adequate social skills that might be of pivotal importance for future challenges like successful group-living and securing competitive fitness advantages.     

The social functions of babbling: acoustic and contextual characteristics that facilitate maternal responsiveness

What is the social function of babbling? An important function of prelinguistic vocalizing may be to elicit parental behavior in ways that facilitate the infant's own learning about speech and language. Infants use parental feedback to their babbling to learn new vocal forms, but the microstructure of parental responses to babbling has not been studied. To enable precise manipulation of the proximal infant cues that may influence maternal behavior, we used a playback paradigm to assess mothers’ responsiveness to prerecorded audiovisual clips of unfamiliar infants’ noncry prelinguistic vocalizations and actions. Acoustic characteristics and directedness of vocalizations were manipulated to test their efficacy in structuring social interactions. We also compared maternal responsiveness in the playback paradigm and in free play with their own infants. Maternal patterns of reactions to babbling were stable across both tasks. In the playback task, we found specific vocal cues, such as the degree of resonance and the transition timing of consonant‐vowel syllables, predicted contingent maternal responding. Vocalizations directed at objects also facilitated increased responsiveness. The responses mothers exhibited, such as sensitive speech and vocal imitation, are known to facilitate vocal learning and development. Infants, by influencing the behavior of their caregivers with their babbling, create social interactions that facilitate their own communicative development.     

Should I stay or should I go? Initiation of joint travel in mother–infant dyads of two chimpanzee communities in the wild

It is well established that great apes communicate via intentionally produced, elaborate and flexible gestural means. Yet relatively little is known about the most fundamental steps into this communicative endeavour—communicative exchanges of mother–infant dyads and gestural acquisition; perhaps because the majority of studies concerned captive groups and single communities in the wild only. Here, we report the first systematic, quantitative comparison of communicative interactions of mother–infant dyads in two communities of wild chimpanzees by focusing on a single communicative function: initiation of carries for joint travel. Over 156 days of observation, we recorded 442 actions, 599 cases of intentional gesture production, 51 multi-modal combinations and 80 vocalisations in the Kanyawara community, Kibale National Park, Uganda, and the Taï South community, Taï National Park, Côte d’Ivoire. Our results showed that (1) mothers and infants differed concerning the signal frequency and modality employed to initiate joint travel, (2) concordance rates of mothers’ gestural production were relatively low within but also between communities, (3) infant communicative development is characterised by a shift from mainly vocal to gestural means, and (4) chimpanzee mothers adjusted their signals to the communicative level of their infants. Since neither genetic channelling nor ontogenetic ritualization explains our results satisfactorily, we propose a revised theory of gestural acquisition, social negotiation, in which gestures are the output of social shaping, shared understanding and mutual construction in real time by both interactants.     

Aping expressions? Chimpanzees produce distinct laugh types when responding to laughter of others

Humans have the ability to replicate the emotional expressions of others even when they undergo different emotions. Such distinct responses of expressions, especially positive expressions, play a central role in everyday social communication of humans and may give the responding individuals important advantages in cooperation and communication. The present work examined laughter in chimpanzees to test whether nonhuman primates also use their expressions in such distinct ways. The approach was first to examine the form and occurrence of laugh replications (laughter after the laughter of others) and spontaneous laughter of chimpanzees during social play and then to test whether their laugh replications represented laugh-elicited laugh responses (laughter triggered by the laughter of others) by using a quantitative method designed to measure responses in natural social settings. The results of this study indicated that chimpanzees produce laugh-elicited laughter that is distinct in form and occurrence from their spontaneous laughter. These findings provide the first empirical evidence that nonhuman primates have the ability to replicate the expressions of others by producing expressions that differ in their underlying emotions and social implications. The data further showed that the laugh-elicited laugh responses of the subjects were closely linked to play maintenance, suggesting that chimpanzees might gain important cooperative and communicative advantages by responding with laughter to the laughter of their social partners. Notably, some chimpanzee groups of this study responded more with laughter than others, an outcome that provides empirical support of a socialization of expressions in great apes similar to that of humans.     

Theory‐of‐Mind development and early sibling relationships after the birth of a sibling: Parental discipline matters

This study investigated relations among children's Theory‐of‐Mind (ToM) development, early sibling interactions, and parental discipline strategies during the transition to siblinghood. Using a sample of firstborn children and their parents (N = 208), we assessed children's ToM before the birth of a sibling and 12 months after the birth, and sibling interactions (i.e., positive engagement and antagonism) and parental discipline strategies (i.e., child‐centred and parent‐centred discipline) at 4 and 8 months in the first year of siblinghood. Structural equation modelling analyses revealed that children's ToM before the birth of the sibling predicted children's positive engagement with the infant sibling, whereas children's antagonistic behaviours towards the infant sibling negatively predicted children's ToM at 12 months, but only when mothers used low levels of child‐centred discipline. These findings emphasize the role of parents in the development of young children's social‐cognitive understanding in the context of early sibling interactions.

Highlights

• This study investigated relations among firstborns' Theory‐of‐Mind (ToM), early sibling relationships, and parental discipline during the first year of siblinghood.
• Multigroup analyses showed that ToM predicted higher sibling positive engagement, and early sibling antagonism predicted poorer ToM when mothers used low child‐centred discipline.
• Parental discipline plays an important role in the development of young children's social understanding and sibling relationships as early as the first year of siblinghood.

     

Roost-mate communication in adult Indian false vampire bats (Megaderma lyra): an indication of individuality in temporal and spectral pattern

The remarkable cognitive abilities of bats indicate that they may recognise particular conspecifics. Because of their highly developed auditory system, it is obvious that vocalisations of bats may give information about the individual emitting them. In a field study of the social behaviour in the Indian false vampire bat (Megaderma lyra), two different types of vocalisation were recorded and analysed. The bats emitted these vocalisations only while aggregating with conspecifics inside the day roost. The “landing strophe” consisted of a number of brief multiharmonic downward frequency-modulated (FM_down) sounds which levelled off as a constant frequency (CF), and the “clatter strophe” was composed of a number of multiharmonic FM_down sounds which became shallow at the end as a short CF. The sounds of the landing strophe and the ¶clatter strophe differed in repetition rate, duration, harmonic components and frequency. Time pattern and peak frequency of the two sound types differed highly significantly between single, unidentified bats. The sounds were inter-individually distinct when the three parameters were combined as an acoustical space. Therefore, these vocalisations might be used for individual recognition in adult bats.     

More than one kind of happiness: Can we recognize vocal expressions of different positive states?

Several theorists have proposed that distinctions are needed between different positive emotional states, and that these discriminations may be particularly useful in the domain of vocal signals (Ekman, 1992b, Cognition and Emotion, 6, 169–200; Scherer, 1986, Psychological Bulletin, 99, 143–165). We report an investigation into the hypothesis that positive basic emotions have distinct vocal expressions (Ekman, 1992b, Cognition and Emotion, 6, 169–200). Non-verbal vocalisations are used that map onto five putative positive emotions: Achievement/Triumph, Amusement, Contentment, Sensual Pleasure, and Relief. Data from categorisation and rating tasks indicate that each vocal expression is accurately categorised and consistently rated as expressing the intended emotion. This pattern is replicated across two language groups. These data, we conclude, provide evidence for the existence of robustly recognisable expressions of distinct positive emotions.     

Spanish-English bilingual toddlers’ vocabulary skills: The role of caregiver language input and warmth

There is a well-documented link between bilingual language development and the relative amounts of exposure to each language. Less is known about the role of quality indicators of caregiver-child interactions in bilingual homes, including caregiver input diversity, warmth and sensitivity. This longitudinal study examines the relation between caregiver input (lexical diversity, amount), warmth and sensitivity and bilingual toddlers’ subsequent vocabulary outcomes. We video-recorded caregiver-child interactions in Spanish-English Latino homes when toddlers (n = 47) were 18 months of age (M = 18.32 months; SD = 1.02 months). At the 24-month follow-up, we measured children's vocabulary as total vocabulary (English, Spanish combined) as well as within language (Spanish, English). Results revealed that Spanish lexical diversity exposure at 18 months from caregivers was positively associated with children's Spanish and total vocabulary scores at 24 months, while English lexical diversity was positively associated with children's English scores; lexical diversity and amount were highly correlated. Additionally, caregivers’ warmth was positively associated with children's Spanish, English and total vocabulary scores. Together, these factors accounted for substantial variance (30–40%) in vocabulary outcomes. Notably, caregiver input accounted for more variance in single language outcomes than did caregiver warmth, whereas caregiver warmth uniquely accounted for more variance in total vocabulary scores. Our findings extend prior research findings by suggesting that children's dual language development may depend on their exposure to a diverse set of words, not only amount of language exposure, as well as warm interactions with caregivers. A video abstract of this article can be viewed at https://youtu.be/q1V_7fz5wog

Highlights

• Video-recorded observations of caregiver-child interactions revealed warmth and high sensitivity from Latino caregivers.
• Linguistically-detailed analyses of caregiver input revealed wide variation in the diversity of Spanish and English directed at 18-month-old bilingual toddlers.
• Bilingual toddlers’ vocabulary (single language, total) was positively associated with caregivers’ diverse input and warmth, thus extending prior findings on bilinguals’ amount of language exposure.
• Findings suggest that caregivers’ lexical diversity explains more variance in bilingual toddlers’ single language outcomes, whereas warmth explains more variance in total vocabulary scores.

     

Children over-imitate adults and peers more than puppets

Researchers commonly use puppets in development science. Amongst other things, puppets are employed to reduce social hierarchies between child participants and adult experimenters akin to peer interactions. However, it remains controversial whether children treat puppets like real-world social partners in these settings. This study investigated children's imitation of causally irrelevant actions (i.e., over-imitation) performed by puppet, adult, or child models. Seventy-two German children (Age_Range = 4.6–6.5 years; 36 girls) from urban, socioeconomically diverse backgrounds observed a model retrieving stickers from reward containers. The model performed causally irrelevant actions either in contact with the reward container or not. Children were more likely to over-imitate adults’ and peers’ actions as compared to puppets’ actions. Across models, they copied contact actions more than no-contact actions. While children imitate causally irrelevant actions from puppet models to some extent, their social learning from puppets does not necessarily match their social learning from real-world social agents, such as children or adults.

Research Highlights

• We examined children's over-imitation from adult, child, and puppet models to validate puppetry as an approach to simulate non-hierarchical interactions.
• Children imitated adults and child models at slightly higher rates than puppets.
• This effect was present regardless of whether the irrelevant actions involved physical contact to the reward container or not.
• In our study children's social learning from puppets does not match their social learning from human models.

     

Learning to pause: Fidelity of and biases in the developmental acquisition of gaps in the communicative signals of a songbird

The temporal organization of sounds used in social contexts can provide information about signal function and evoke varying responses in listeners (receivers). For example, music is a universal and learned human behavior that is characterized by different rhythms and tempos that can evoke disparate responses in listeners. Similarly, birdsong is a social behavior in songbirds that is learned during critical periods in development and used to evoke physiological and behavioral responses in receivers. Recent investigations have begun to reveal the breadth of universal patterns in birdsong and their similarities to common patterns in speech and music, but relatively little is known about the degree to which biological predispositions and developmental experiences interact to shape the temporal patterning of birdsong. Here, we investigated how biological predispositions modulate the acquisition and production of an important temporal feature of birdsong, namely the duration of silent pauses (“gaps”) between vocal elements (“syllables”). Through analyses of semi-naturally raised and experimentally tutored zebra finches, we observed that juvenile zebra finches imitate the durations of the silent gaps in their tutor's song. Further, when juveniles were experimentally tutored with stimuli containing a wide range of gap durations, we observed biases in the prevalence and stereotypy of gap durations. Together, these studies demonstrate how biological predispositions and developmental experiences differently affect distinct temporal features of birdsong and highlight similarities in developmental plasticity across birdsong, speech, and music.

Research Highlights

• The temporal organization of learned acoustic patterns can be similar across human cultures and across species, suggesting biological predispositions in acquisition.
• We studied how biological predispositions and developmental experiences affect an important temporal feature of birdsong, namely the duration of silent intervals between vocal elements (“gaps”).
• Semi-naturally and experimentally tutored zebra finches imitated the durations of gaps in their tutor's song and displayed some biases in the learning and production of gap durations and in gap variability.
• These findings in the zebra finch provide parallels with the acquisition of temporal features of speech and music in humans.

     

Children's associations between space and pitch are differentially shaped by language

Musical properties, such as auditory pitch, are not expressed in the same way across cultures. In some languages, pitch is expressed in terms of spatial height (high vs. low), whereas others rely on thickness vocabulary (thick = low frequency vs. thin = high frequency). We investigated how children represent pitch in the face of this variable linguistic input by examining the developmental trajectory of linguistic and non-linguistic space-pitch associations in children who acquire Dutch (a height-pitch language) or Turkish (a thickness-pitch language). Five-year-olds, 7-year-olds, 9-year-olds, and 11-year-olds were tested for their understanding of pitch terminology and their associations of spatial dimensions with auditory pitch when no language was used. Across tasks, thickness-pitch associations were more robust than height-pitch associations. This was true for Turkish children, and also Dutch children not exposed to thickness-pitch vocabulary. Height-pitch associations, on the other hand, were not reliable—not even in Dutch-speaking children until age 11—the age when they demonstrated full comprehension of height-pitch terminology. Moreover, Turkish-speaking children reversed height-pitch associations. Taken together, these findings suggest thickness-pitch associations are acquired in similar ways by children from different cultures, but the acquisition of height-pitch associations is more susceptible to linguistic input. Overall, then, despite cross-cultural stability in some components, there is variation in how children come to represent musical pitch, one of the building blocks of music.

Research Highlights

• Children from diverse cultures differ in their understanding of music vocabulary and in their nonlinguistic associations between spatial dimensions and auditory pitch.
• Height-pitch mappings are acquired late and require additional scaffolding from language, whereas thickness-pitch mappings are acquired early and are less susceptible to language input.
• Space-pitch mappings are not static from birth to adulthood, but change over development, suggesting music cognition is shaped by cross-cultural experience.

     

Chimpanzee Use of Human and Conspecific Social Cues to Locate Hidden Food

Two studies are reported in which chimpanzees attempted to use social cues to locate hidden food in one of two possible hiding places. In the first study four chimpanzees were exposed to a local enhancement cue (the informant approached and looked to the location where food was hidden and then remained beside it) and a gaze/point cue (the informant gazed and manually pointed towards the location where the food was hidden). Each cue was given by both a human informant and a chimpanzee informant. In the second study 12 chimpanzees were exposed to a gaze direction cue in combination with a vocal cue (the human informant gazed to the hiding location and produced one of two different vocalizations: a ‘food-bark’ or a human word-form). The results were: (i) all subjects were quite skillful with the local enhancement cue, no matter who produced it; (ii) few subjects were skillful with the gaze/point cue, no matter who produced it (most of these being individuals who had been raised in infancy by humans); and (iii) most subjects were skillful when the human gazed and vocalized at the hiding place, with little difference between the two types of vocal cue. Findings are discussed in terms of chimpanzees’ apparent need for additional cues, over and above gaze direction cues, to indicate the presence of food.     

The development of representational play in chimpanzees and bonobos: Evolutionary implications,pretense, and the role of interspecies communication

This research investigates the development of symbolic or representational play in two species of the genus Pan, bonobos (Pan paniscus) and chimpanzees (Pan troglodytes). The participants varied not only by species, but also as to whether they had become proficient in communicating with humans via a set of arbitrary visual symbols, called lexigrams. Using a developmental sequence of representational play based on McCune, we found every level that children manifest to be constructed by Pan. The most robust and regular ontogenetic sequence for both bonobos and chimpanzee was not McCune's five-level progression, but a three-step ontogenetic sequence: Level 1 (no representation, no pretense) precedes Levels 2–4 (representation but no pretense), which in turn precedes Level 5 (includes pretense as well as representation). A linguistic system for interspecies communication was necessary for Level 5 representational play and “true” pretense. Human scaffolding produced developmental progress within sequences for all the apes, except the bonobo who lacked a system of interspecies communication. This evidence suggests that the potential for representational play and its social stimulation were present in the common ancestor of bonobos, chimpanzees and humans five million years ago.     

The repertoire and intentionality of gestural communication in wild chimpanzees

A growing body of evidence suggests that human language may have emerged primarily in the gestural rather than vocal domain, and that studying gestural communication in great apes is crucial to understanding language evolution. Although manual and bodily gestures are considered distinct at a neural level, there has been very limited consideration of potential differences at a behavioural level. In this study, we conducted naturalistic observations of adult wild East African chimpanzees (Pan troglodytes schweinfurthii) in order to establish a repertoire of gestures, and examine intentionality of gesture production, use and comprehension, comparing across manual and bodily gestures. At the population level, 120 distinct gesture types were identified, consisting of 65 manual gestures and 55 bodily gestures. Both bodily and manual gestures were used intentionally and effectively to attain specific goals, by signallers who were sensitive to recipient attention. However, manual gestures differed from bodily gestures in terms of communicative persistence, indicating a qualitatively different form of behavioural flexibility in achieving goals. Both repertoire size and frequency of manual gesturing were more affiliative than bodily gestures, while bodily gestures were more antagonistic. These results indicate that manual gestures may have played a significant role in the emergence of increased flexibility in great ape communication and social bonding.     

Temporal Relationships Between Gaze and Vocal Behavior in Prelinguistic and Linguistic Communication

This work reports longitudinal evaluation of the temporal relationships between gaze and vocal behavior addressed to interactive partners (mother or experimenter) in a free-play situation. Thirteen children were observed at the ages of 1;0 and 1;8 during laboratory sessions, and video recordings of free-play interactions with mother and a female experimenter were coded separately for children's vocal behavior (vocalizations and words) and gaze toward their interactive partners. The difference between the observed and expected cooccurrence of these two communicative behaviors was evaluated by transformation into z-scores. The most important findings are related to differences in the temporal relationship observed at age 1;0 between gaze and vocalizations and at age 1;8 between gaze and words. At the earlier age, the infants who exhibited greater coordination between gaze and vocal behavior than was expected by chance (z-score > +1.96) preferred to look at the interlocutor at the beginning of the vocal turn. Instead, when they were older and began to produce words, they frequently looked at the interlocutor at the end of the vocal turn. These results are interpreted as referring to characteristics of conversational competence in the prelinguistic and linguistic periods. Moreover, looking at the interlocutor at the beginning of the vocal turn at age 1;0 was found to be related to language production at age 1:8, highlighting a significant relationship between conversational competence during the prelinguistic period and language acquisition.