Tool use by chimpanzees (Pan troglodytes) of the Arnhem Zoo community

Abstract: Chimpanzees have a large repertoire of tool-use behaviors. This study reports on the variety and the extent of tool use exhibited by the chimpanzees of the Arnhem Zoo community in The Netherlands, living in an enriched captive setting since 1971. The use of tools by 29 chimpanzees aged from 0 to 37 years was observed. We identified 13 types of tool use comparable to those found in the wild. Some of these types of tool use seem to be specific to this community, and can be explained by the ecological characteristics of this captive setting. Chimpanzees started to use tools from the age of 2 years. Young chimpanzees, from 5 to 9 years old, showed a greater repertoire of tool use than infants and adults. All types of tool use in the community have appeared by the age of 10, the age of puberty for chimpanzees. Multivariate analysis was applied for the 29 individuals by 13 types of tool use in a one-zero matrix. The results show two major categories of tool use, one in a practical or substantial context and the other in a nonpractical or play context. The subjects clustered into groups reflecting developmental stages, although there are great individual differences. In conclusion, this captive community provides a unique opportunity to clarify the details of tool use by chimpanzees.     

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.     

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.     

Skill mastery inhibits adoption of observed alternative solutions among chimpanzees (Pan troglodytes)

Geographic variation in socially transmitted skills and signals, similar to human culture, has been well documented for great apes. The rules governing the adoption of novel behaviours, however, are still largely unknown. We conducted an innovation-and-transmission experiment with two groups of chimpanzees living at hopE Primate Sanctuary Gänserndorf, Austria, presenting a board on which food had to be manoeuvred around obstacles to be acquired. Most chimpanzees used sticks to acquire the food, but five adults independently invented a novel technique, rattling, which was subsequently tested by almost all group members. However, individuals who had become proficient with sticks were reluctant to switch to rattling, despite it being more efficient. Similarly, after rattling was prevented, rattle specialists kept trying to rattle and made no attempt to use the stick technique, despite their knowledge about its existence. We conclude that innovators stimulate others to experiment with the solutions they display, but that chimpanzees are nevertheless conservative; mastery of a skill inhibits further exploration, and hence adoption of alternative techniques even if these are more efficient. Consequently, conformity among group members should not be expected in great apes when individuals develop proficiency at different techniques. Conservatism thus joins conformity as a mechanism to bring about cultural uniformity and stability.     

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.     

Differing views: Can chimpanzees do Level 2 perspective-taking?

Although chimpanzees understand what others may see, it is unclear whether they understand how others see things (Level 2 perspective-taking). We investigated whether chimpanzees can predict the behavior of a conspecific which is holding a mistaken perspective that differs from their own. The subject competed with a conspecific over two food sticks. While the subject could see that both were the same size, to the competitor one appeared bigger than the other. In a previously established game, the competitor chose one stick in private first and the subject chose thereafter, without knowing which of the sticks was gone. Chimpanzees and 6-year-old children chose the ‘riskier’ stick (that looked bigger to the competitor) significantly less in the game than in a nonsocial control. Children chose randomly in the control, thus showing Level 2 perspective-taking skills; in contrast, chimpanzees had a preference for the ‘riskier’ stick here, rendering it possible that they attributed their own preference to the competitor to predict her choice. We thus run a follow-up in which chimpanzees did not have a preference in the control. Now, they also chose randomly in the game. We conclude that chimpanzees solved the task by attributing their own preference to the other, while children truly understood the other’s mistaken perspective.     

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.     

Antisocial Behavior,Moral Disengagement,Empathy and Negative Emotion: A Comparison Between Disabled and Able-Bodied Athletes

Theories of morality suggest that negative emotions associated with antisocial behavior should diminish motivation for such behavior. Two reasons that have been proposed to explain why some individuals repeatedly harm others are that (a) they use mechanisms of moral disengagement to justify their actions, and (b) they may not empathize with and vicariously experience the negative emotions felt by their victims. With the aim of testing these proposals, the present study compared spinal cord injured disabled athletes and able-bodied athletes to determine the effect of reduced visceral afferent feedback caused by spinal cord injury on antisocial behavior, moral disengagement, empathy, and negative emotion (i.e., anger, anxiety, and dejection). Disabled athletes reported less frequent antisocial behavior and lower moral disengagement than able-bodied athletes. Group differences in antisocial behavior were mediated by differences in moral disengagement. The two groups did not differ in empathy or negative emotion. The findings of this study suggest that antisocial behavior may be regulated by mechanisms of moral disengagement.     

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.     

Wild chimpanzees (Pan troglodytes schweinfurthii) distinguish between different scream types: evidence from a playback study

When experiencing aggression from group members, chimpanzees commonly produce screams. These agonistic screams are graded signals and vary acoustically as a function of the severity of aggression the caller is facing. We conducted a series of field playback experiments with a community of wild chimpanzees in the Budongo Forest, Uganda, to determine whether individuals could meaningfully distinguish between screams given in different agonistic contexts. We compared six subjects’ responses to screams given in response to severe and mild aggression. Subjects consistently discriminated between the two scream types. To address the possibility that the response differences were driven directly by the screams’ peripheral acoustic features, rather than any attached social meaning, we also tested the subjects’ responses to tantrum screams. These screams are given by individuals that experienced social frustration, but no physical threat, yet acoustically they are very similar to screams of victims of severe aggression. We found chimpanzees looked longer at severe victim screams than either mild victim screams or tantrum screams. Our results indicate that chimpanzees attend to the informational content of screams and are able to distinguish between different scream variants, which form part of a graded continuum.     

Attention Deficit Disorder and hospitalization due to injury among older adolescents in New South Wales,Australia

The association between Attention Deficit Disorder (ADD) and injuries among children has long been studied. More recently, research interest has extended to the long-term consequences of childhood ADD in adolescents and young adults, mainly focusing on motor vehicle-related accidents, but not other injuries. Very little work has investigated the associations between individual characteristics, type of injuries, outcomes, and the diagnosis of ADD upon admission of injured patients to a hospital. To investigate these associations, this study used a population-based epidemiological design that analyzed data routinely collected on patients hospitalized due to injuries. Results showed that younger, male, and more socioeconomically disadvantaged patients were more likely to be diagnosed with ADD upon admission. Significant association between different causes of injuries, in particular self-inflicted injuries, and diagnosis of ADD were also found. Patients diagnosed with ADD stayed in the hospital longer than others disregarding the cause of their injury. These results illustrate clearly that ADD is underrecognized among injured patients upon admission to hospitals. Information on the ADD status of injured patients is important for treatment decision making. A screening tool should be developed for assessing all injured patients upon admission to hospitals.     

Heritability of hand preference in chimpanzees (Pan troglodytes): evidence from a partial interspecies cross-fostering study.

Heritability of hand preference was tested in a sample of 188 chimpanzees (Pan troglodytes). Hand preference was measured by coordinated bimanual actions, and concordance percentages were compared between parents and offspring and siblings. Among siblings, concordance percentages were compared for dyads in which both individuals were raised by chimpanzees, both were raised by humans, or 1 was raised in each environment. The results indicated population-level right hand preferences for coordinated bimanual actions. There were no significant associations in hand preference between parents and offspring. In full and maternal half siblings, concordance in hand preference was significantly greater than chance in mother- and human-reared individuals but not in cross-fostered dyads. The cumulative results suggest that the direction of hand preference is heritable in chimpanzees but the mechanism of transmission is not genetic. Several environmental explanations are proposed to explain the findings, including the potential role of maternal cradling bias and in utero fetal position.     

Body-Region-Specific Injuries as Predictors of Psychosocial Outcomes Among Those Injured in Combat: Results From the Wounded Warrior Recovery Project

Associations between body region injured and psychosocial outcomes may have implications for injury prevention and mitigation strategies. The present study investigated the association of body-region—specific injuries and their association with 3 psychosocial outcomes (i.e., quality of life, QOL; posttraumatic stress disorder, PTSD; and depression) among a large sample of U.S. military service members injured in combat. A total of 1,011 individuals wounded in combat enrolled in the Wounded Warrior Recovery Project and provided QOL, depression, and PTSD measures on a Web-based survey. These psychosocial variables were linked with the presence/absence of combat injuries in 9 different body regions (head, face, neck, thorax, spine, abdomen, upper extremity, lower extremity, and external skin/other), as well as overall injury severity and demographic factors. Two-step hierarchical linear regressions revealed that after adjusting for overall injury severity and time since injury, those with combat-related head and spine injuries were particularly at risk for relatively worse psychosocial outcomes. Head and spinal injuries were associated with lower QOL and higher PTSD and depression. Overall, even those with relatively minor injuries may be at risk of lower QOL and adverse psychosocial outcomes. These findings may highlight the need for early interventions to minimize the psychological effects of combat injuries to the head and spine.     

Components of analogical reasoning in a mildly head injured population

This study sought to investigate the effects of mild head injury on a particular type of cognitive ability, verbal analogical reasoning. The performance of 19 individuals with head injuries was compared to a group of 30 control subjects matched for age, education, and gender on 100 verbal analogies. Solution times and error rates were modeled. Unstandardized regression weights for individual subjects were correlated with subjects’ performance on a number of standardized ability tests. Results showed that compared to the control subjects, the head injured subjects: (a) were significantly slower to solve the analogies, and were particularly slow to perform certain processes: encoding/inference and comparison; (b) tended to show greater variability in performance; and (c) had data that had a poorer componential model fit. The data suggest that analogical reasoning is affected by a head injury, and that certain information processes may be responsible for performance deficits.     

Gender differences in susceptibility to posttraumatic stress disorder

We examined data from a community survey of trauma exposure and DSM-IV PTSD in Winnipeg, Canada to explore factors associated with the higher rate of PTSD in women than men. Women were found to be at significantly increased risk for PTSD following exposure to serious trauma (odds ratio approximately 5), even when sexual trauma--which predominates in women--was excluded (odds ratio approximately 3). Adjusting for gender differences in the number of lifetime traumata, or in the likelihood of the trauma being associated with particular reactions to or consequences of the event (i.e. thinking that one would be killed or seriously injured; sustaining a serious physical injury; seeing someone else seriously injured or killed) did not result in a lessening of the PTSD risk in women. Women were found to be at increased risk for PTSD following nonsexual assaultive violence (e.g. mugging or other physical attack) but not following non-assaultive trauma (e.g. fire, witnessing injury to others). Understanding the basis of (and parameters for) this increased susceptibility to PTSD in women compared to men following trauma exposure should be a priority for future traumatic stress research.     

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.

     

Great apes' understanding of other individuals' line of sight

Previous research has shown that many social animals follow the gaze of other individuals. However, knowledge about how this skill differs between species and whether it shows a relationship with genetic distance from humans is still fragmentary. In the present study of gaze following in great apes, we manipulated the nature of a visual obstruction and the presence/absence of a target. We found that bonobos, chimpanzees, and gorillas followed gaze significantly more often when the obstruction had a window than when it did not, just as human infants do. Additionally, bonobos and chimpanzees looked at the experimenter's side of a windowless obstruction more often than the other species. Moreover, bonobos produced more double looks when the barrier was opaque than when it had a window, indicating an understanding of what other individuals see. The most distant human relatives studied, orangutans, showed few signs of understanding what another individual saw. Instead, they were attracted to the target's location by the target's presence, but not by the experimenter's gaze. Great apes' perspective-taking skills seem to have increased in the evolutionary lineage leading to bonobos, chimpanzees, and humans.     

Chimpanzees prioritise social information over pre-existing behaviours in a group context but not in dyads

How animal communities arrive at homogeneous behavioural preferences is a central question for studies of cultural evolution. Here, we investigated whether chimpanzees (Pan troglodytes) would relinquish a pre-existing behaviour to adopt an alternative demonstrated by an overwhelming majority of group mates; in other words, whether chimpanzees behave in a conformist manner. In each of five groups of chimpanzees (N?=?37), one individual was trained on one method of opening a two-action puzzle box to obtain food, while the remaining individuals learned the alternative method. Over 5 h of open access to the apparatus in a group context, it was found that 4/5 ‘minority’ individuals explored the majority method and three of these used this new method in the majority of trials. Those that switched did so after observing only a small subset of their group, thereby not matching conventional definitions of conformity. In a further ‘Dyad’ condition, six pairs of chimpanzees were trained on alternative methods and then given access to the task together. Only one of these individuals ever switched method. The number of observations that individuals in the minority and Dyad individuals made of their untrained method was not found to influence whether or not they themselves switched to use it. In a final ‘Asocial’ condition, individuals (N?=?10) did not receive social information and did not deviate from their first-learned method. We argue that these results demonstrate an important influence of social context upon prioritisation of social information over pre-existing methods, which can result in group homogeneity of behaviour.     

Manual dexterity in the gorilla: bimanual and digit role differentiation in a natural task

The manipulative actions of mountain gorillas Gorilla g. beringei were examined in the context of foraging on hard-to-process plant foods in the field, in particular those used in tackling thistle Carduus nyassanus. A repertoire of 72 functionally distinct manipulative actions was recorded. Many of these actions were used in several variants of grip, finger(s) and movement path, both by different individuals and by the same individual at different times. The repertoire appears somewhat greater than that observed in comparable studies of monkeys, but a far more striking difference is found in the use of differentiated actions in concert. Mountain gorillas routinely and frequently deal with problems that involve: (1) bimanual role differentiation, with the two hands taking different roles but synchronized in time and space, and (2) digit role differentiation, with independent control of parts of the same hand used for separate purposes at the same time. The independent control that allows these abilities, so crucial to human manual constructional ability, is apparently general in African great apes. Role differentiation, between and within the hand, is evidently a primitive characteristic in the human arsenal of skills. Accepted after revision: 6 April 2001 Electronic Publication