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.     

Referential communication by chimpanzees (Pan troglodytes)

Two experiments were conducted to assess the referential function of chimpanzee (Pan troglodytes) gestures to obtain food. The chimpanzees received 1 trial per condition. In Experiment 1 (N = 101), in full view of the chimpanzee, a banana was placed on top of 1 of 2 inverted buckets or was hidden underneath 1 of the buckets. In Experiment 2 (N = 35), 4 conditions were presented in constant order: (a) no food, no observer; (b) no food, observer present; (c) food present, no observer; and (d) food present, observer present. Gestures and visual orienting were used socially and referentially. The capacity for nonverbal reference may predate the Hominidae-Pongidae split, and the development of nonverbal reference may be independent of human species-specific adaptations for speech.     

Scale-model comprehension by chimpanzees (Pan troglodytes)

The ability of chimpanzees (Pan troglodytes) to recognize the correspondence between a scale model and its real-world referent was examined. In Experiments 1 and 2, an adult female and a young adult male watched as an experimenter hid a miniature model food in 1 of 4 sites in a scale model. Then, the chimpanzees were given the opportunity to find the real food item that had been hidden in the analogous location in the real room. The female performed significantly above chance, whereas the male performed at chance level. Experiments 3 and 4 tested 5 adult and 2 adolescent chimpanzees in a similar paradigm, using a scale model of the chimpanzees' outdoor area. Results indicate that some adult chimpanzees were able to reliably demonstrate an understanding of the relationship between a scale model and the larger space it represented, whereas other subjects were constrained by inefficient and unsuccessful search patterns.     

Factors influencing imitation of manipulatory actions in chimpanzees (Pan troglodytes).

The purpose of the study was to investigate what kind of factors determine the degree of difficulty for chimpanzees (Pan troglodytes) when they imitate actions. Five adult chimpanzees were instructed to perform 48 arbitrary manipulatory actions consisting of different bodily motor patterns and object directionality. Results showed that actions in which an object is directed toward another external location (another object and one's own body) were easier to perform than those that involved manipulating a single object alone. Actions involving unfamiliar motor patterns were more difficult to perform than those involving familiar motor patterns that were already present in the subject's repertoire. Error responses were characterized as perseverative repetition of previously instructed actions. These findings suggest that chimpanzees find the directionality of manipulated objects a more salient cue than details of the demonstrator's body movements performing the manipulation.     

Multimodal communication by captive chimpanzees (Pan troglodytes)

Many studies have shown that apes and monkeys are adept at cross-modal matching tasks requiring the subject to identify objects in one modality when information regarding those objects has been presented in a different modality. However, much less is known about non-human primates’ production of multimodal signaling in communicative contexts. Here, we present evidence from a study of 110 chimpanzees demonstrating that they select the modality of communication in accordance with variations in the attentional focus of a human interactant, which is consistent with previous research. In each trial, we presented desirable food to one of two chimpanzees, turning mid-way through the trial from facing one chimpanzee to facing the other chimpanzee, and documented their communicative displays, as the experimenter turned towards or away from the subjects. These chimpanzees varied their signals within a context-appropriate modality, displaying a range of different visual signals when a human experimenter was facing them and a range of different auditory or tactile (attention-getting) signals when the human was facing away from them; this finding extends previous research on multimodal signaling in this species. Thus, in the impoverished circumstances characteristic of captivity, complex signaling tactics are nevertheless exhibited by chimpanzees, suggesting continuity in intersubjective psychological processes in humans and apes.     

Spontaneous spatial constructions by chimpanzees (Pan troglodytes,Pan paniscus)

We investigated chimpanzees’ spontaneous spatial constructions with objects. Two were common chimpanzees ages 6 and 18 years and one was an 11‐year‐old bonobo raised in a very enriched human environment from very early in life. Chimpanzees’ products and procedures were quite advanced when constructing functional spatial relations by placing objects in or on each other, but not when constructing nonfunctional spatial relations by placing objects next to each other. This difference was found in all their constructions, including their spatial correspondences and spatial symmetries. Compared to those of younger nonencultured chimpanzees all their spatial constructions were more advanced. Compared to human infants, their functional spatial constructions were also more advanced but their nonfunctional spatial constructions were less advanced.     

Imitation in neonatal chimpanzees (Pan troglodytes)

This paper provides evidence for imitative abilities in neonatal chimpanzees (Pan troglodytes), our closest relatives. Two chimpanzees were reared from birth by their biological mothers. At less than 7 days of age the chimpanzees could discriminate between, and imitate, human facial gestures (tongue protrusion and mouth opening). By the time they were 2 months old, however, the chimpanzees no longer imitated the gestures. They began to perform mouth opening frequently in response to any of the three facial gestures presented to them. These findings suggest that neonatal facial imitation is most likely an innate ability, developed through natural selection in humans and in chimpanzees. The relationship between the disappearance of neonatal imitation and the development of social communicative behavior is discussed from an evolutionary perspective.     

Summation and numerousness judgments of sequentially presented sets of items by chimpanzees (Pan troglodytes).

Summation and numerousness judgments by 2 chimpanzees (Pan troglodytes) were investigated when 2 quantities of M&Ms were presented sequentially, and the quantities were never viewed in their totality. Each M&M was visible only before placement in 1 of 2 cups. In Experiment 1, sets of 1 to 9 M&Ms were presented. In Experiment 2, the quantities in each cup were presented as 2 smaller sets (e.g., 2 + 2 vs. 4 + 1). In Experiment 3, the quantities were presented as 3 smaller sets (e.g., 2 + 2 + 3 vs. 3 + 4 + 1). In Experiment 4, an M&M was removed from 1 set before the chimpanzees' selection. In Experiments 1 and 2, the chimpanzees selected the larger quantity on significantly more trials than would be predicted by chance. In Experiments 3 and 4, 1 chimpanzee performed at a level significantly better than chance. Therefore, chimpanzees mentally represent quantity and successfully combine and compare nonvisible, sequentially presented sets of items.     

Spontaneous discrimination of natural stimuli by chimpanzees (Pan troglodytes)

Six chimpanzees (Pan troglodytes) were presented with pairs of color photographic images of 5 different categories of animals (cat, chimp, gorilla, tiger, fish). The subjects responded to each pair using symbols for "same" and "different." Both within- and between-category discriminations were tested, and all chimpanzees classified the image pairs in accordance with the 5 experimenter-defined categories under conditions of nondifferential reinforcement. Although previous studies have demonstrated identification or discrimination of natural categories by nonhuman animals, subjects were typically differentially reinforced for their responses. The present findings demonstrate that chimpanzees can classify natural objects spontaneously and that such classifications may be similar to those that would be observed in human subjects.     

Spontaneous transfer of matching by infant chimpanzees (Pan troglodytes)

Four infant chimpanzees learned a matching-to-sample task when only two training stimuli were used. They then spontaneously transferred the matching concept to novel items, including three-dimensional objects and fabric swatches, without any experimenter-provided differential feedback. These results support the view that the matching concept is broadly construed by chimpanzees from the beginning and does not depend upon explicit training.     

Spontaneous classification in action by a human-enculturated and language-reared bonobo (Pan paniscus) and common chimpanzees (Pan troglodytes).

The authors investigated the formal features of spontaneous manipulations used by 1 bonobo (Pan paniscus) and 2 common chimpanzees (Pan troglodytes) to classify objects in action. Chimpanzees' manipulations were evenly split between serial, one-at-a-time acts on 1 object and parallel, two-at-a-time acts on 2 or 3 objects. Chimpanzees systematically combined their manipulations into routines to generate class-consistent categories of objects. Their routines featured much reproduction of the same manipulations, planful acts that anticipated follow-up manipulations, and manipulations that were reciprocal to each other to accomplish an end. In all these respects, chimpanzees' manipulations were similar to those of 2-year-old human infants. In others they differed. Chimpanzees' routines were mainly based on a linear integration of manipulations. Classifying in action was only infrequently produced by hierarchically integrated routines.     

Inferences about guessing and knowing by chimpanzees (Pan troglodytes)

The visual perspective-taking ability of 4 chimpanzees (Pan troglodytes) was investigated. The subjects chose between information about the location of hidden food provided by 2 experimenters who randomly alternated between two roles (the guesser and the knower). The knower baited 1 of 4 obscured cups so that the subjects could watch the process but could not see which of the cups contained the reward. The guesser waited outside the room until the food was hidden. Finally, the knower pointed to the correct cup while the guesser pointed to an incorrect one. The chimpanzees quickly learned to respond to the knower. They also showed transfer to a novel variation of the task, in which the guesser remained inside the room and covered his head while the knower stood next to him and watched a third experimenter bait the cups. The results are consistent with the hypothesis that chimpanzees are capable of modeling the visual perspectives of others.     

Constructive and deconstructive tool modification by chimpanzees (Pan troglodytes)

Nine chimpanzees (Pan troglodytes) were tested for their ability to assemble or disassemble the appropriate tool to obtain a food reward from two different apparatus. In its deconstructed form, the tool functioned as a probe for one apparatus. In its constructed form, the tool functioned as a hook, appropriate for a second apparatus. Each subject completed four test trials with each apparatus type. Tool types were randomized and counter-balanced between the two forms. Results demonstrated that adult and juvenile chimpanzees (N = 7) were successful with both tool types, while two infant chimpanzees performed near chance. Off-line video analyses revealed that tool modifications followed by attempted solutions by the adults and juveniles were typically correct on the first attempt. Neither infant was successful in modifying tools correctly on the first attempt over all eight trials. The older chimpanzees’ ability to modify the appropriate tool consistently prior to use indicates an immediate recognition of the functional attributes necessary for the successful use of tool types on each apparatus, and represents a non-replication of a previously reported study by Povinelli.

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.     

Lateral bias in infant chimpanzees (Pan troglodytes)

This study documents the presence, strength, and direction of lateralization in chimpanzees (Pan troglodytes) over the first 3 months of life. Nursery-reared chimpanzees (7 males and 5 females) were repeatedly assessed on a behavioral scale. Lateral bias was measured for 4 behaviors: hand-to-mouth, hand-to-hand, defensive grasp, and first step. Hand-to-mouth was significantly lateralized for the sample. Eight of the 10 chimpanzees that showed hand-to-mouth used the right hand. Lateral bias for defensive grasp was positively related to lateral bias both of first step and of hand-to-mouth. Lateral bias in hand-to-mouth was inversely related to lateral bias in hand-to-hand. Strength of lateralization increased as chimpanzees matured. These laterality effects in infant chimpanzees were expressed under conditions of emotional arousal. Moreover, degree of laterality may be a predictor of responsivity to stress.     

Interactive use of sign language by cross-fostered chimpanzees (Pan troglodytes)

Cross-fostered as infants in Reno, Nevada, chimpanzees (Pan troglodytes) Washoe, Moja, Tatu, and Dar freely converse in signs of American Sign Language with each other as well as with humans in Ellensburg, Washington. In this experiment, a human interlocutor waited for a chimpanzee to initiate conversations with her and then responded with 1 of 4 types of probes: general requests for more information, on-topic questions, off-topic questions, or negative statements. The responses of the chimpanzees to the probes depended on the type of probe and the particular signs in the probes. They reiterated, adjusted, and shifted the signs in their utterances in conversationally appropriate rejoinders. Their reactions to and interactions with a conversational partner resembled patterns of conversation found in similar studies of human children.     

Strategies used to combine seriated cups by chimpanzees (Pan troglodytes), bonobos (Pan paniscus), and capuchins (Cebus apella).

The authors investigated strategies used to combine seriated cups by apes (Pan troglodytes and P. paniscus) and monkeys (Cebus apella) using a protocol reported in P. M. Greenfield, K. Nelson, and E. Saltzman's (1972) study with children. It was hypothesized that apes would exhibit more hierarchical combinations of cups than monkeys, given apes' language capacity, and that apes would seriate the cups more efficiently than monkeys. As predicted, apes made many structures with the cups using a variety of strategies, and monkeys rarely combined the cups. After a training phase to orient monkeys to the task, the 2 genera did not differ in the strategies used to combine the cups or in efficiency in seriating the cups. Success in this task suggests that sensorimotor versions of hierarchically organized combinatorial activity are well within apes' and monkeys' abilities.     

Numerical judgments by chimpanzees (Pan troglodytes) in a token economy

We presented four chimpanzees with a series of tasks that involved comparing two token sets or comparing a token set to a quantity of food. Selected tokens could be exchanged for food items on a one-to-one basis. Chimpanzees successfully selected the larger numerical set for comparisons of 1 to 5 items when both sets were visible and when sets were presented through one-by-one addition of tokens into two opaque containers. Two of four chimpanzees used the number of tokens and food items to guide responding in all conditions, rather than relying on token color, size, total amount, or duration of set presentation. These results demonstrate that judgments of simultaneous and sequential sets of stimuli are made by some chimpanzees on the basis of the numerousness of sets rather than other non-numerical dimensions. The tokens were treated as equivalent to food items on the basis of their numerousness, and the chimpanzees maximized reward by choosing the larger number of items in all situations.     

A group-specific arbitrary tradition in chimpanzees (Pan troglodytes)

Social learning in chimpanzees has been studied extensively and it is now widely accepted that chimpanzees have the capacity to learn from conspecifics through a multitude of mechanisms. Very few studies, however, have documented the existence of spontaneously emerged traditions in chimpanzee communities. While the rigour of experimental studies is helpful to investigate social learning mechanisms, documentation of naturally occurring traditions is necessary to understand the relevance of social learning in the real lives of animals. In this study, we report on chimpanzees spontaneously copying a seemingly non-adaptive behaviour (“grass-in-ear behaviour”). The behaviour entailed chimpanzees selecting a stiff, straw-like blade of grass, inserting the grass into one of their own ears, adjusting the position, and then leaving it in their ear during subsequent activities. Using a daily focal follow procedure, over the course of 1 year, we observed 8 (out of 12) group members engaging in this peculiar behaviour. Importantly, in the three neighbouring groups of chimpanzees (n = 82), this behaviour was only observed once, indicating that ecological factors were not determiners of the prevalence of this behaviour. These observations show that chimpanzees have a tendency to copy each other’s behaviour, even when the adaptive value of the behaviour is presumably absent.     

Quantity judgments of auditory and visual stimuli by chimpanzees (Pan troglodytes)

Many species can choose between two visual sets of stimuli on the basis of quantity. This is true when sets are both visible, or are presented one set at a time or even one item at a time. However, we know comparatively little about how well nonhuman animals can compare auditory quantities. Here, three chimpanzees (Pan troglodytes) chose between two sets of food items when they only heard each item fall into different containers rather than seeing those items. This method prevented the chimpanzees from summing the amount of visible food they saw because there were no visual cues. Chimpanzees performed well, and their performance matched that of previous experiments with regard to obeying Weber's law. They also performed well with comparisons between a sequentially presented auditory set and a fully visible set, demonstrating that duration of presentation was not being used as a cue. In addition, they accommodated empty sets into these judgments, although not perfectly. Thus, chimpanzees can judge auditory quantities in flexible ways that show many similarities to how they compare visual quantities.