The distribution and development of handedness for manual gestures in captive chimpanzees (Pan troglodytes)

This article describes the distribution and development of handedness for manual gestures in captive chimpanzees. Data on handedness for unimanual gestures were collected in a sample of 227 captive chimpanzees. Handedness for these gestures was compared with handedness for three other measures of hand use: tool use, reaching, and coordinated bimanual actions. Chimpanzees were significantly more right-handed for gestures than for all other measures of hand use. Hand use for simple reaching at 3 to 4 years of age predicted hand use for gestures 10 years later. Use of the right hand for gestures was significantly higher when gestures were accompanied by a vocalization than when they were not. The collective results suggest that left-hemisphere specialization for language may have evolved initially from asymmetries in manual gestures in the common ancestor of chimpanzees and humans, rather than from hand use associated with other, non-communicative motor actions, including tool use and coordinated bimanual actions, as has been previously suggested in the literature.     

Cultural innovation and transmission of tool use in wild chimpanzees: evidence from field experiments

Chimpanzees (Pan troglodytes) are the most proficient and versatile users of tools in the wild. How such skills become integrated into the behavioural repertoire of wild chimpanzee communities is investigated here by drawing together evidence from three complementary approaches in a group of oil-palm nut- (Elaeis guineensis) cracking chimpanzees at Bossou, Guinea. First, extensive surveys of communities adjacent to Bossou have shown that population-specific details of tool use, such as the selection of species of nuts as targets for cracking, cannot be explained purely on the basis of ecological differences. Second, a 16-year longitudinal record tracing the development of nut-cracking in individual chimpanzees has highlighted the importance of a critical period for learning (3–5 years of age), while the similar learning contexts experienced by siblings have been found to result in near-perfect (13 out of 14 dyads) inter-sibling correspondence in laterality. Third, novel data from field experiments involving the introduction of unfamiliar species of nuts to the Bossou group illuminates key aspects of both cultural innovation and transmission. We show that responses of individuals toward the novel items differ markedly with age, with juveniles being the most likely to explore. Furthermore, subjects are highly specific in their selection of conspecifics as models for observation, attending to the nut-cracking activities of individuals in the same age group or older, but not younger than themselves. Together with the phenomenon of inter-community migration, these results demonstrate a mechanism for the emergence of culture in wild chimpanzees.     

Cognitive capacities for behavioural flexibility in wild chimpanzees (Pan troglodytes): the effect of snare injury on complex manual food processing

In chimpanzees, it is only in the restricted context of tool use that manual and cognitive skills have been described, comparable to those that gorillas and orang-utans display in obtaining plant foods. We report the complex food preparation skills used to eat, without tools, the leaves of the tree Broussonettia papyrifera in the Sonso community of chimpanzees at Budongo Forest, Uganda. Able-bodied individuals used multi-stage techniques that required bimanual role differentiation at several stages, and were hierarchical in organisation. A total repertoire of 14 techniques was found, with strong preference in all individuals for either of two of these; 6 additional techniques were found when flowers and leaves were eaten together. However, in this community over 20% of individuals suffer from some form of upper- or lower-limb injury as a result of snares. We investigated the manner of compensation for upper-limb injury. Only the most severely injured showed reduced feeding efficiency. Injured individuals were found to use the same repertoire of techniques as able-bodied chimpanzees. We found no evidence to suggest that injured individuals were able to develop wholly novel techniques optimal for their specific injuries, although shifts in preference for particular techniques did occur. Rather, injured individuals used novel ways of achieving particular steps in the process; by "working around" their impairments; in this way, they managed to use the same techniques as the able-bodied. Since snare injuries generally befall young animals, these results suggest that chimpanzees learn techniques partly through observational learning (of, necessarily, able-bodied individuals). Accepted after revision: 8 February 2001 Electronic Publication     

Vocal learning of a communicative signal in captive chimpanzees, Pan troglodytes

We hypothesized that chimpanzees could learn to produce attention-getting (AG) sounds via positive reinforcement. We conducted a vocal assessment in 76 captive chimpanzees for their use of AG sounds to acquire the attention of an otherwise inattentive human. Fourteen individuals that did not produce AG sounds during the vocal assessment were evaluated for their ability to acquire the use of an AG sound through operant conditioning and to employ these sounds in an attention-getting context. Nine of the 14 chimpanzees were successfully shaped using positive reinforcement to produce an AG sound. In a post-training vocal assessment, eight of the nine individuals that were successfully trained to produce AG sounds generalized the use of these newly acquired signals to communicatively relevant situations. Chimpanzees possess the ability to acquire the use of a communicative signal via operant conditioning and can generalize the use of this newly acquired signal to appropriate communicative contexts.     

Chimpanzee (Pan troglodytes) intentional communication is not contingent upon food

Studies of great apes have revealed that they use manual gestures and other signals to communicate about distal objects. There is also evidence that chimpanzees modify the types of communicative signals they use depending on the attentional state of a human communicative partner. The majority of previous studies have involved chimpanzees requesting food items from a human experimenter. Here, these same communicative behaviors are reported in chimpanzees requesting a tool from a human observer. In this study, captive chimpanzees were found to gesture, vocalize, and display more often when the experimenter had a tool than when she did not. It was also found that chimpanzees responded differentially based on the attentional state of a human experimenter, and when given the wrong tool persisted in their communicative efforts. Implications for the referential and intentional nature of chimpanzee communicative signaling are discussed.     

Spontaneous class grouping behavior by bonobos (Pan paniscus) and common chimpanzees (P. troglodytes)

Two experiments investigated spontaneous class grouping behavior by human-enculturated and language-reared bonobos (Pan paniscus) and common chimpanzees (P. troglodytes). In experiment 1, three chimpanzees ranging in age from 6 to 18 years were presented with six objects. The objects embodied three conditions: additive, multiplicative and disjoint classes. All chimpanzees spontaneously produced single- and two-category classifying. In experiment 2, six chimpanzees ranging in age from 6 to 21 years were presented with 12 objects in the same class conditions. Chimpanzees mainly produced single-category classifying. Their two-category classifying was more rudimentary than that found in experiment 1. Chimpanzees did not produce any three-category classifying which would be necessary to construct the hierarchies that humans begin to construct during early childhood. Received: 1 February 1999 / Accepted after revision: 9 August 1999     

Spontaneous logicomathematical constructions by chimpanzees (Pan troglodytes, P. paniscus)

Two experiments investigated the spontaneous construction of precursory logicomathematical operations by human-enculturated and language-reared chimpanzees (Pan troglodytes, Pan paniscus) when they were interacting freely with objects. In experiment 1, three chimpanzees ranging in age from 6 to 18 years were presented with sets of six objects. Chimpanzees constructed equivalence, order and reversibility relations within single sets of objects as well as between two or three contemporaneous sets of objects. The chimpanzees’ logicomathematical operations were more advanced, including infrequent and minimal operations on three sets, than those of some previously investigated younger nonenculturated common chimpanzees. In experiment 2, six chimpanzees ranging in age from 6 to 21 years were presented with sets of 12 objects. Chimpanzees constructed more advanced operations on single sets, but not on contemporaneous sets. The results suggest partial convergence and partial divergence between development of logicomathematical cognition in chimpanzees and humans. Received: 20 January 1999 / Accepted after revision: 9 August 1999     

The gestural repertoire of the wild chimpanzee

Great ape gestural communication is known to be intentional, elaborate and flexible; yet there is controversy over the best interpretation of the system and how gestures are acquired, perhaps because most studies have been made in restricted, captive settings. Here, we report the first systematic analysis of gesture in a population of wild chimpanzees. Over 266 days of observation, we recorded 4,397 cases of intentional gesture use in the Sonso community, Budongo, Uganda. We describe 66 distinct gesture types: this estimate appears close to asymptote, and the Sonso repertoire includes most gestures described informally at other sites. Differences in repertoire were noted between individuals and age classes, but in both cases, the measured repertoire size was predicted by the time subjects were observed gesturing. No idiosyncratic usages were found, i.e. no gesture type was used only by one individual. No support was found for the idea that gestures are acquired by ‘ontogenetic ritualization’ from originally effective actions; moreover, in detailed analyses of two gestures, action elements composing the gestures did not closely match those of the presumed original actions. Rather, chimpanzee gestures are species-typical; indeed, many are ‘family-typical’, because gesture types recorded in gorillas, orangutans and chimpanzee overlap extensively, with 24 gestures recorded in all three genera. Nevertheless, chimpanzee gestures are used flexibly across a range of contexts and show clear adjustment to audience (e.g. silent gestures for attentive targets, contact gestures for inattentive ones). Such highly intentional use of a species-typical repertoire raises intriguing questions for the evolution of advanced communication.     

Differential use of vocal and gestural communication by chimpanzees (Pan troglodytes) in response to the attentional status of a human (Homo sapiens)

This study examined the communicative behavior of 49 captive chimpanzees (Pan troglodytes), particularly their use of vocalizations, manual gestures, and other auditory- or tactile-based behaviors as a means of gaining an inattentive audience's attention. A human (Homo sapiens) experimenter held a banana while oriented either toward or away from the chimpanzee. The chimpanzees' behavior was recorded for 60 s. Chimpanzees emitted vocalizations faster and were more likely to produce vocalizations as their 1st communicative behavior when a human was oriented away from them. Chimpanzees used manual gestures more frequently and faster when the human was oriented toward them. These results replicate the findings of earlier studies on chimpanzee gestural communication and provide new information about the intentional and functional use of their vocalizations.     

Raking it in: the impact of enculturation on chimpanzee tool use

Recent evidence for different tool kits, proposed to be based upon culture-like transmission, have been observed across different chimpanzee communities across Western Africa. In light of these findings, the reported failures by seven captive juvenile chimpanzees tested with 27 tool use tasks (Povinelli 2000) seem enigmatic. Here we report successful performance by a group of nine captive, enculturated chimpanzees, and limited success by a group of six semi-enculturated chimpanzees, on two of the Povinelli tasks, the Flimsy Tool task, and the Hybrid Tool task. All chimpanzees were presented with a rake with a flimsy head and a second rake with a rigid head, either of which could be used to attempt to retrieve a food reward that was out of reach. The rigid rake was constructed such that it had the necessary functional features to permit successful retrieval, while the flimsy rake did not. Both chimpanzee groups in the present experiment selected the functional rigid tool correctly to use during the Flimsy Tool task. All animals were then presented with two “hybrid rakes” A and B, with one half of each rake head constructed from flimsy, non-functional fabric, and the other half of the head was made of wood. Food rewards were placed in front of the rigid side of Rake A and the flimsy side of Rake B. To be successful, the chimps needed to choose the rake that had the reward in front of the rigid side of the rake head. The fully enculturated animals were successful in selecting the functional rake, while the semi-enculturated subjects chose randomly between the two hybrid tools. Compared with findings from Povinelli, whose non-enculturated animals failed both tasks, our results demonstrate that chimpanzees reared under conditions of semi-enculturation could learn to discriminate correctly the necessary tool through trial-and-error during the Flimsy Tool task, but were unable to recognize the functional relationship necessary for retrieving the reward with the “hybrid” rake. In contrast, the enculturated chimpanzees were correct in their choices during both the Flimsy Tool and the Hybrid Tool tasks. These results provide the first empirical evidence for the differential effects of enculturation on subsequent tool use capacities in captive chimpanzees.     

How to crack nuts: acquisition process in captive chimpanzees (Pan troglodytes) observing a model

Stone tool use for nut cracking consists of placing a hard-shelled nut onto a stone anvil and then cracking the shell open by pounding it with a stone hammer to get to the kernel. We investigated the acquisition of tool use for nut cracking in a group of captive chimpanzees to clarify what kind of understanding of the tools and actions will lead to the acquisition of this type of tool use in the presence of a skilled model. A human experimenter trained a male chimpanzee until he mastered the use of a hammer and anvil stone to crack open macadamia nuts. He was then put in a nut-cracking situation together with his group mates, who were naïve to this tool use; we did not have a control group without a model. The results showed that the process of acquisition could be broken down into several steps, including recognition of applying pressure to the nut, emergence of the use of a combination of three objects, emergence of the hitting action, using a tool for hitting, and hitting the nut. The chimpanzees recognized these different components separately and practiced them one after another. They gradually united these factors in their behavior leading to their first success. Their behavior did not clearly improve immediately after observing successful nut cracking by a peer, but observation of a skilled group member seemed to have a gradual, long-term influence on the acquisition of nut cracking by naïve chimpanzees.     

Tool-composite reuse in wild chimpanzees (Pan troglodytes): archaeologically invisible steps in the technological evolution of early hominins?

Recent etho-archaeological studies of stone-tool use by wild chimpanzees have contributed valuable data towards elucidating the variables that influenced the emergence and development of the first lithic industries among Plio-Pleistocene hominins. Such data help to identify potential behaviours entailed in the first percussive technologies that are invisible in archaeological records. The long-term research site of Bossou in Guinea features a unique chimpanzee community whose members systematically use portable stones as hammers and anvils to crack open nuts in natural as well as in field experimental settings. Here we present the first analysis of repeated reuse of the same tool-composites in wild chimpanzees. Data collected over 5 years of experimental nut-cracking sessions at an “outdoor laboratory” site were assessed for the existence of systematic patterns in the selection of tool-composites, at group and at individual levels. Chimpanzees combined certain stones as hammer and anvil more often than expected by chance, even when taking into account preferences for individual stones by themselves. This may reflect an ability to recognise the nut-cracker as a single tool (composed of two elements, but functional only as a whole), as well as discrimination of tool quality-effectiveness. Through repeatedly combining the same pairs of stones—whether due to preferences for particular composites or for the two elements independently—tool-users may amplify use-wear traces and increase the likelihood of fracturing the stones, and thus of detaching pieces by battering.     

Serial gesturing by wild chimpanzees: its nature and function for communication

Chimpanzees at Budongo, Uganda, regularly gesture in series, including ‘bouts’ of gesturing that include response waiting and ‘sequences’ of rapid-fire gesturing without pauses. We examined the distribution and correlates of 723 sequences and 504 bouts for clues to the function of multigesture series. Gesturing by older chimpanzees was more likely to be successful, but the success rate of any particular gesture did not vary with signaller age. Rather, older individuals were more likely to choose successful gestures, and these highly successful gestures were more often used singly. These patterns explain why bouts were recorded most in younger animals, whereas older chimpanzees relied more on single gestures: bouts are best interpreted as a consequence of persistence in the face of failure. When at least one gesture of a successful type occurred in a sequence, that sequence was more likely to be successful; overall, however, sequences were less successful than single gestures. We suggest that young chimpanzees use sequences as a ‘fail-safe’ strategy: because they have the innate potential to produce a large and redundant repertoire of gestures but lack knowledge of which of them would be most efficient. Using sequences increases the chance of giving one effective gesture and also allows users to learn the most effective types. As they do so, they need to use sequences less; sequences may remain important for subtle interpersonal adjustment, especially in play. This ‘Repertoire Tuning’ hypothesis explains a number of results previously reported from chimpanzee gesturing.     

Acquisition and Transmission of Tool Making and Use for Drinking Juice in a Group of Captive Chimpanzees (Pan troglodytes)

The present study examined the acquisition and transmission of tool making and use in a group of chimpanzees. We set up a piece of apparatus that provided orange juice in an outdoor compound for a group of nine chimpanzees. Although they could reach the juice with their hands, eight of the nine subjects used tools. Fifteen kinds of tools in total were used, such as straw, twigs, and some kinds of leaves. The chimpanzees showed high selectivity with regard to tool type. They preferred to use Thuja occidentalis as a tool although there were 28 species of tree and several kinds of grass available in the compound. Two females initiated the use of the Thuja tool. Since then, five other individuals have begun to use it selectively. Before making the tools by themselves, these five chimpanzees first watched others using the Thuja tool for drinking juice, and then used the Thuja tool which had been used and left by another chimpanzee.     

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.

     

Development of self-recognition in chimpanzees (Pan troglodytes).

Chimpanzees (Pan troglodytes) demonstrate the ability to recognize themselves in mirrors, yet investigations of the development of self-recognition in chimpanzees are sparse. Twelve young chimpanzees, grouped by age, were given mirror exposure and tested for self-recognition and contingent movement. All 6 juveniles, 4 and 5 years old, exhibited mirror-guided, mark-directed behavior and clear evidence of self-recognition. In contrast, among the infants, only the oldest group of 2 1/2-year-olds exhibited clear evidence of self-recognition. All chimpanzees exhibited both self-directed behaviors and contingent movements. These results suggest that self-recognition occurs at a slightly older age in chimpanzees than in human infants. In humans, self-recognition is linked with other cognitive abilities. The results conform to the general pattern that great apes exhibit many cognitive skills comparable to those of 2-year-old humans.     

The gestural repertoire of the wild bonobo (<Emphasis Type="Italic">Pan paniscus</Emphasis>): a mutually understood communication system

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

Now you see me,now you don't: evidence that chimpanzees understand the role of the eyes in attention

Chimpanzees appear to understand something about the attentional states of others; in the present experiment, we investigated whether they understand that the attentional state of a human is based on eye gaze. In all, 116 adult chimpanzees were offered food by an experimenter who engaged in one of the four experimental manipulations: eyes closed, eyes open, hand over eyes, and hand over mouth. The communicative behavior of the chimpanzees was observed. More visible behaviors were produced when the experimenter's eyes were visible than when the experimenter's eyes were not visible. More vocalizations were produced when the experimenter's eyes were closed than when they were open, but there were no differences in other attention getting behaviors. There was no effect of age or rearing history. The results suggest that chimpanzees use the presence of the eyes as a cue that their visual gestures will be effective.