Forest chimpanzees (Pan troglodytes verus) remember the location of numerous fruit trees

It is assumed that spatial memory contributes crucially to animal cognition since animals’ habitats entail a large number of dispersed and unpredictable food sources. Spatial memory has been investigated under controlled conditions, with different species showing and different conditions leading to varying performance levels. However, the number of food sources investigated is very low compared to what exists under natural conditions, where food resources are so abundant that it is difficult to precisely identify what is available. By using a detailed botanical map containing over 12,499 trees known to be used by the Taï chimpanzees, we created virtual maps of all productive fruit trees to simulate potential strategies used by wild chimpanzees to reach resources without spatial memory. First, we simulated different assumptions concerning the chimpanzees’ preference for a particular tree species, and, second, we varied the detection field to control for the possible use of smell to detect fruiting trees. For all these assumptions, we compared simulated distance travelled, frequencies of trees visited, and revisit rates with what we actually observed in wild chimpanzees. Our results show that chimpanzees visit rare tree species more frequently, travel shorter distances to reach them, and revisit the same trees more often than if they had no spatial memory. In addition, we demonstrate that chimpanzees travel longer distances to reach resources where they will eat for longer periods of time, and revisit resources more frequently where they ate for a long period of time during their first visit. Therefore, this study shows that forest chimpanzees possess a precise spatial memory which allows them to remember the location of numerous resources and use this information to select the most attractive resources.     

Social influences on ant-dipping acquisition in the wild chimpanzees (Pan troglodytes verus) of Bossou,Guinea, West Africa

We currently have little understanding of the influence of learning opportunity, whether social or environmental, and maternal role on tool-use acquisition in young wild chimpanzees. This study aims to fill this gap by focusing on the acquisition of ant-dipping among chimpanzees of Bossou, Guinea. Ant-dipping is a hazardous tool-use behaviour aimed at army ants (Dorylus spp.). Bossou chimpanzees target these ants both at nests (high risk) and trails (low risk) and employ two techniques to consume them: direct mouthing and pull-through. We present data for 13 mother–offspring pairs (1–10 years old). Mothers with young ≤5 years old dipped significantly more often at trails than at nests, thus minimizing the risk posed to themselves and their young. Infants thus benefited from better conditions to observe and practice ant-dipping. Mothers also varied greatly in their percent time spent ant-dipping and offspring differed in their learning opportunity. Our results suggest that high opportunity young started to observe and perform ant-dipping sooner and were better at ant-dipping than low opportunity young. Although mothers and weaned offspring correlated positively in their percent time spent dipping and proficiency, they did not match in technique used or tool length. Finally, we propose that the learning trajectory of young may predict individual and sex differences in adulthood. This study demonstrates the important role of mothers and learning opportunity in the acquisition of a hazardous tool-use behaviour and suggests that chimpanzee material culture is a product of a complex interaction between social processes and ecological factors.     

Dominance assertion in male chimpanzees (Pan troglodytes)

The social relations of three adult males in a captive chimpanzee group were evaluated for 12 months. The observations encompassed a period of transition in the dominance hierarchy providing an opportunity to assess behavioral patterns involved in the initiation of status changes. This study shows that dominance in the chimpanzee is expressed primarily through ritualized and non-contact behavior rather than by overt aggression. In particular, display behavior is important for indicating dominance rank, in communicating aggressive intent, and as a prelude to status struggles. The functional relationships between status, displays and other socio-sexual behavior patterns are presented.     

Overlooked evidence for semantic compositionality and signal reduction in wild chimpanzees (Pan troglodytes)

Animal Cognition - Recent discoveries of semantic compositionality in Japanese tits have enlivened the discussions on the presence of this phenomenon in wild animal communication. Data on semantic...     

Development of face recognition in infant chimpanzees (Pan troglodytes)

In this paper, we assessed the developmental changes in face recognition by three infant chimpanzees aged 1–18 weeks, using preferential-looking procedures that measured the infants’ eye- and head-tracking of moving stimuli. In Experiment 1, we prepared photographs of the mother of each infant and an “average” chimpanzee face using computer-graphics technology. Prior to 4 weeks of age, the infants showed few tracking responses and no differential responses. Between 4 and 8 weeks of age, they paid greater attention to their mother's face. From 8 weeks onward, they again showed no differences, but exhibited frequent tracking responses. Experiment 2 investigated the infants’ tracking responses between a familiar human's and an “average” human face. The infants did not show any evidence of recognizing the human faces. We discuss the development of face recognition in relation to the effects of other species’ faces and postnatal visual experience.     

Imitation of intentional manipulatory actions in chimpanzees (Pan troglodytes)

In this study, the authors investigated the understanding of other's actions in 5 adult chimpanzees (Pan troglodytes). A human demonstrated an attempt to open different containers. Each container required a different motor pattern to open it. Along with the container, a 2nd object was made available. After a free play period in which the chimpanzees' natural behaviors toward the objects were recorded, the authors tested the following 2 phases: The demonstrator (a) tried but failed to open and (b) opened the container successfully, with 1 of 2 alternative strategies, either using an "irrelevant tool" or by hand. The chimpanzees did not reproduce the demonstrator's motor patterns precisely but did reproduce the demonstrated strategies in both phases. These results suggest that chimpanzees anticipate the intentions of others by perceiving the directionality and causality of object(s) as available cues.     

Agonistic screams in wild chimpanzees (Pan troglodytes schweinfurthii) vary as a function of social role

Some nonhuman primates have demonstrated the capacity to communicate about external objects or events, suggesting primate vocalizations can function as referential signals. However, there is little convincing evidence for functionally referential communication in any great ape species. Here, the authors demonstrate that wild chimpanzees (Pan troglodytes schweinfurthii) of Budongo forest, Uganda, give acoustically distinct screams during agonistic interactions depending on the role they play in a conflict. The authors analyzed the acoustic structure of screams of 14 individuals, in the role of both aggressor and victim. The authors found consistent differences in the acoustic structure of the screams, across individuals, depending on the social role the individual played during the conflict. The authors propose that these 2 distinct scream variants, produced by victims and aggressors during agonistic interactions, may be promising candidates for functioning as referential signals.     

The generalization of deferred imitation in enculturated chimpanzees (Pan troglodytes)

Deferred imitation of object-related actions and generalization of imitation to similar but not identical tasks was assessed in three human-reared (enculturated) chimpanzees, ranging in age from 5 to 9 years. Each ape displayed high levels of deferred imitation and only slightly lower levels of generalization of imitation. The youngest two chimpanzees were more apt to generalize the model's actions when they had displayed portions of the target behaviors at baseline, consistent with the idea that learning is more likely to occur when working within the "zone of proximal development." We argue that generalization of imitation is the best evidence to date of imitative learning in chimpanzees. Electronic Publication     

Group differences in the mutual gaze of chimpanzees (Pan troglodytes)

A comparative developmental framework was used to determine whether mutual gaze is unique to humans and, if not, whether common mechanisms support the development of mutual gaze in chimpanzees and humans. Mother-infant chimpanzees engaged in approximately 17 instances of mutual gaze per hour. Mutual gaze occurred in positive, nonagonistic contexts. Mother-infant chimpanzees at a Japanese center exhibited significantly more mutual gaze than those at a center in the United States. Cradling and motor stimulation varied across groups. Time spent cradling infants was inversely related to mutual gaze. It is suggested that in primates, mutual engagement is supported via an interchangeability of tactile and visual modalities. The importance of mutual gaze is best understood within a perspective that embraces both cross-species and cross-cultural data.     

Preference for human direct gaze in infant chimpanzees (Pan troglodytes)

We studied gaze perception in three infant chimpanzees (Pan troglodytes), aged 10-32 weeks, using a two-choice preferential-looking paradigm. The infants were presented with two photographs of a human face: (a) with the eyes open or closed, and (b) with a direct or an averted gaze. We found that the chimpanzees preferred looking at the direct-gaze face. However, in the context of scrambled faces, the infants showed no difference in gaze discrimination between direct and averted gazes. These findings suggest that gaze perception by chimpanzees may be influenced by the surrounding facial context. The relationship between gaze perception, face processing, and the adaptive significance of gaze perception are discussed from an evolutionary perspective.     

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.     

Spatial memory and monitoring of hidden items through spatial displacements by chimpanzees (Pan troglodytes)

This study examined chimpanzee (Pan troglodytes) short-term memory for food location in near space. In Experiments 1 and 2, either 1 or 2 items (chocolate pieces) were hidden in an array of 3 or 5 containers that either remained stationary or were rotated 180 degrees or 360 degrees. When the array remained stationary, the chimpanzees remembered both item locations. When arrays were rotated, however, chimpanzees found only 1 item. In Experiment 3, 2 items were hidden in an array of 7 cups. Both items were found at levels significantly better than chance. Ninety percent of errors were made after the 1st item was found, and errors reflected memory failure rather than a failure of inhibitory control.     

Cognitive and physiological markers of emotional awareness in chimpanzees (Pan troglodytes)

The ability to understand emotion in others is one of the most important factors involved in regulating social interactions in primates. Such emotional awareness functions to coordinate activity among group members, enable the formation of long-lasting individual relationships, and facilitate the pursuit of shared interests. Despite these important evolutionary implications, comparative studies of emotional processing in humans and great apes are practically nonexistent, constituting a major gap in our understanding of the extent to which emotional awareness has played an important role in shaping human behavior and societies. This paper presents the results of two experiments that examine chimpanzees' responses to emotional stimuli. First, changes in peripheral skin temperature were measured while subjects viewed three categories of emotionally negative video scenes; conspecifics being injected with needles (INJ), darts and needles alone (DART), and conspecific directing agonism towards the veterinarians (CHASE). Second, chimpanzees were required to use facial expressions to categorize emotional video scenes, i.e., favorite food and objects and veterinarian procedures, according to their positive and negative valence. With no prior training, subjects spontaneously matched the emotional videos to conspecific facial expressions according to their shared emotional meaning, indicating that chimpanzee facial expressions are processed emotionally, as are human expressions. Decreases in peripheral skin temperature, indicative of negative sympathetic arousal, were significantly lower when subjects viewed the INJ and DART videos, compared to the CHASE videos, indicating greater negative arousal when viewing conspecifics being injected with needles, and needles themselves, than when viewing conspecifics engaged in general agonism. Accepted after revision: 8 April 2001 ❚ Electronic Publication     

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.     

Perception of emergent configurations in humans (Homo sapiens) and chimpanzees (Pan troglodytes)

We examined the perceptions of emergent configurations in humans and chimpanzees using a target-localization task. The stimulus display consisted of a target placed among multiple identical distractors. The target and distractors were presented either solely, within congruent contexts in which salient configurations emerge, or within incongruent contexts in which salient configurations do not emerge. We found that congruent contexts had similar facilitative effects on target localization by humans and chimpanzees, whereas similar disruptive effects emerged when the stimuli were presented within incongruent contexts. When display size was manipulated, targets under the congruent-context condition were localized in a parallel manner, but those under the no-context and incongruent-context conditions were localized in a serial manner by both species. These results suggest that both humans and chimpanzees perceive emergent configurations when targets and distractors are presented within certain congruent contexts and that they process such emergent configurations preattentively.     

Emergence of symmetry in a visual conditional discrimination by chimpanzees (Pan troglodytes)

In Exp. 1, three young chimpanzees were trained to match red to a cross and green to a circle in an arbitrary matching-to-sample task. After acquisition of this task, they were tested for the emergence of associative symmetry of these conditional relations using the trials on which shapes were presented as samples and colors as comparisons. One of the three chimpanzees showed statistically significant accuracy on these test trials. This successful subject served in Exp. 2, in which an auditory-visual stimulus appeared contingent upon red and a cross while another auditory-visual stimulus was contingent upon green and a circle. This subject showed higher accuracies in symmetry tests than in Exp. 1, which suggested the facilitative effects of these events common to sample and correct comparison on the development of symmetry. In Exp. 3, subsequent tests in which only these stimuli were presented as samples indicated that these stimuli might have become the members of equivalence classes.     

Acquisition of navigation by chimpanzees (Pan troglodytes) in an automated fingermaze task

These experiments investigated how chimpanzees learn to navigate visual fingermazes presented on a touch monitor. The aim was to determine whether training the subjects to solve several different mazes would establish a generalized map-reading skill such that they would solve new mazes correctly on the first presentation. In experiment 1, two captive adult female chimpanzees were trained to move a visual object (a ball) with a finger over the monitor surface toward a target through a grid of obstacles that formed a maze. The task was fully automated with storage of movement paths on individual trials. Training progressed from very simple mazes with one obstacle to complex mazes with several obstacles. The subjects learned to move the ball to the target in a curved path so as to avoid obstacles and blind alleys. After training on several mazes, both subjects developed a high level of efficiency in moving the ball to the target in a path that closely approached the ideal shortest path. New mazes were then presented to determine whether the subjects had acquired a more generalized maze-solving performance. The subjects solved 65–100% of the new mazes the first time they were presented by moving the ball around obstacles to the target without making detours into blind alleys. In experiment 2, one of the chimpanzees was trained using mazes with two routes to the target. One of the routes was blocked at one of many possible locations. After training to avoid the blind alley in different mazes, new mazes were presented that also had one route blocked. The subject correctly solved 90.7% of the novel mazes. When the mazes had one short and one long open route to the target the subject preferred the shorter route. When the short route was blocked, the subject solved only 53.3% of the mazes because of the preference for the shorter route even when blocked. The overall results suggest that with the training methods used the subjects learned to solve specific mazes with a trial-and-error method. Although both subjects were able to solve many of the novel mazes they did not fully develop a more general "map-reading" skill. Accepted after revision: 30 July 2001 Electronic Publication     

Use of gesture sequences in captive chimpanzee (Pan troglodytes) play

This study examined the use of sensory modalities relative to a partner’s behavior in gesture sequences during captive chimpanzee play at the Chimpanzee and Human Communication Institute. We hypothesized that chimpanzees would use visual gestures toward attentive recipients and auditory/tactile gestures toward inattentive recipients. We also hypothesized that gesture sequences would be more prevalent toward unresponsive rather than responsive recipients. The chimpanzees used significantly more auditory/tactile rather than visual gestures first in sequences with both attentive and inattentive recipients. They rarely used visual gestures toward inattentive recipients. Auditory/tactile gestures were effective with and used with both attentive and inattentive recipients. Recipients responded significantly more to single gestures than to first gestures in sequences. Sequences often indicated that recipients did not respond to initial gestures, whereas effective single gestures made more gestures unnecessary. The chimpanzees thus gestured appropriately relative to a recipient’s behavior and modified their interactions according to contextual social cues.     

Perception of shape from shading in chimpanzees (Pan troglodytes) and humans (Homo sapiens)

The perception of shape from shading was tested in two chimpanzees (Pan troglodytes) and five humans (Homo sapiens), using visual search tasks. Subjects were required to select and touch an odd item (target) from among uniform distractors. Humans found the target faster when shading was vertical than when it was horizontal, consistent with results of previous research. Both chimpanzees showed the opposite pattern: they found the target faster when shading was horizontal. The same difference in response was found in texture segregation tasks. This difference between the species could not be explained by head rotation or head shift parallel to the surface of the monitor. Furthermore, when the shaded shape was changed from a circle to a square, or the shading type was changed from gradual to stepwise, the difference in performance between vertical and horizontal shading disappeared in chimpanzees, but persisted in humans. These results suggest that chimpanzees process shading information in a different way from humans. Received: 20 January 1998 / Accepted after revision: 30 March 1998