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Nielsen, M., Collier-Baker, E., Davis, J. M., & Suddendorf, T. (2005). Imitation recognition in a captive chimpanzee (Pan troglodytes). Anim. Cogn., 8(1), 31–36.
Abstract: This study investigated the ability of a captive chimpanzee (Pan troglodytes) to recognise when he is being imitated. In the experimental condition of test 1a, an experimenter imitated the postures and behaviours of the chimpanzee as they were being displayed. In three control conditions the same experimenter exhibited (1) actions that were contingent on, but different from, the actions of the chimpanzee, (2) actions that were not contingent on, and different from, the actions of the chimpanzee, or (3) no action at all. The chimpanzee showed more “testing” sequences (i.e., systematically varying his actions while oriented to the imitating experimenter) and more repetitive behaviour when he was being imitated, than when he was not. This finding was replicated 4 months later in test 1b. When the experimenter repeated the same actions she displayed in the experimental condition of test 1a back to the chimpanzee in test 2, these actions now did not elicit those same testing sequences or repetitive behaviours. However, a live imitation condition did. Together these results provide the first evidence of imitation recognition in a nonhuman animal.
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Fripp, D., Owen, C., Quintana-Rizzo, E., Shapiro, A., Buckstaff, K., Jankowski, K., et al. (2005). Bottlenose dolphin (Tursiops truncatus) calves appear to model their signature whistles on the signature whistles of community members. Anim. Cogn., 8(1), 17–26.
Abstract: Bottlenose dolphins are unusual among non-human mammals in their ability to learn new sounds. This study investigates the importance of vocal learning in the development of dolphin signature whistles and the influence of social interactions on that process. We used focal animal behavioral follows to observe six calves in Sarasota Bay, Fla., recording their social associations during their first summer, and their signature whistles during their second. The signature whistles of five calves were determined. Using dynamic time warping (DTW) of frequency contours, the calves' signature whistles were compared to the signature whistles of several sets of dolphins: their own associates, the other calves' associates, Tampa Bay dolphins, and captive dolphins. Whistles were considered similar if their DTW similarity score was greater than those of 95% of the whistle comparisons. Association was defined primarily in terms of time within 50 m of the mother/calf pair. On average, there were six dolphins with signature whistles similar to the signature whistles of each of the calves. These were significantly more likely to be Sarasota Bay resident dolphins than non-Sarasota dolphins, and (though not significantly) more likely to be dolphins that were within 50 m of the mother and calf less than 5% of the time. These results suggest that calves may model their signature whistles on the signature whistles of members of their community, possibly community members with whom they associate only rarely.
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Bjorklund, D. F., Yunger, J. L., Bering, J. M., & Ragan, P. (2002). The generalization of deferred imitation in enculturated chimpanzees (Pan troglodytes). Anim. Cogn., 5(1), 49–58.
Abstract: 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.
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Caldwell, C. A., & Whiten, A. (2004). Testing for social learning and imitation in common marmosets, Callithrix jacchus, using an artificial fruit. Anim. Cogn., 7(2), 77–85.
Abstract: We tested for social learning and imitation in common marmosets using an artificial foraging task and trained conspecific demonstrators. We trained a demonstrator marmoset to open an artificial fruit, providing a full demonstration of the task to be learned. Another marmoset provided a partial demonstration, controlling for stimulus enhancement effects, by eating food from the outside of the apparatus. We thus compared three observer groups, each consisting of four animals: those that received the full demonstration, those that received the partial demonstration, and a control group that saw no demonstration prior to testing. Although none of the observer marmosets succeeded in opening the artificial fruit during the test periods, there were clear effects of demonstration type. Those that saw the full demonstration manipulated the apparatus more overall, whereas those from the control group manipulated it the least of the three groups. Those from the full-demonstration group also contacted the particular parts of the artificial fruit that they had seen touched (localised stimulus enhancement) to a greater extent than the other two groups. There was also an interaction between the number of hand and mouth touches made to the artificial fruit for the full- and partial-demonstration groups. Whether or not these data represent evidence for imitation is discussed. We also propose that the clear differences between the groups suggest that social learning mechanisms provide real benefits to these animals in terms of developing novel food-processing skills analogous to the one presented here.
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Bard, K. A. (2007). Neonatal imitation in chimpanzees (Pan troglodytes) tested with two paradigms. Anim. Cogn., 10(2), 233–242.
Abstract: Primate species differ in their imitative performance, perhaps reflecting differences in imitative capacity. The developmentally earliest form of imitation in humans, neonatal imitation, occurs in early interactions with social partners, and may be a more accurate index of innate capacity than imitation of actions on objects, which requires more cognitive ability. This study assessed imitative capacity in five neonatal chimpanzees, within a narrow age range (7-15 days of age), by testing responses to facial and vocal actions with two different test paradigms (structured and communicative). Imitation of mouth opening was found in both paradigms. In the communicative paradigm, significant agreement was found between infant actions and demonstrations. Additionally, chimpanzees matched the sequence of three actions of the TC model, but only on the second demonstration. Newborn chimpanzees matched more modeled actions in the communicative test than in the structured paradigm. These performances of chimpanzees, at birth, are in agreement with the literature, supporting a conclusion that imitative capacity is not unique to the human species. Developmental histories must be more fully considered in the cross-species study of imitation, as there is a greater degree of innate imitative capacity than previously known. Socialization practices interact with innate and developing competencies to determine the outcome of imitation tests later in life.
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Mercado, E. 3rd, Herman, L. M., & Pack, A. A. (2005). Song copying by humpback whales: themes and variations. Anim. Cogn., 8(2), 93–102.
Abstract: Male humpback whales (Megaptera novaeangliae) produce long, structured sequences of sound underwater, commonly called “songs.” Humpbacks progressively modify their songs over time in ways that suggest that individuals are copying song elements that they hear being used by other singers. Little is known about the factors that determine how whales learn from their auditory experiences. Song learning in birds is better understood and appears to be constrained by stable core attributes such as species-specific sound repertoires and song syntax. To clarify whether similar constraints exist for song learning by humpbacks, we analyzed changes over 14 years in the sounds used by humpback whales singing in Hawaiian waters. We found that although the properties of individual sounds within songs are quite variable over time, the overall distribution of certain acoustic features within the repertoire appears to be stable. In particular, our findings suggest that species-specific constraints on temporal features of song sounds determine song form, whereas spectral variability allows whales to flexibly adapt song elements.
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Ducoing, A. M., & Thierry, B. (2005). Tool-use learning in Tonkean macaques (Macaca tonkeana). Anim. Cogn., 8(2), 103–113.
Abstract: The transmission of tool use is a rare event in monkeys. Such an event arose in a group of semi-free-ranging Tonkean macaques (Macaca tonkeana) in which leaning a pole against the park's fence (branch leaning) appeared and spread to several males. This prompted us to test individual and social learning of this behavior in seven young males. In the first experiment, three males learned individually to obtain a food reward using a wooden pole as a climbing tool. They began using the pole to retrieve the reward only when they could alternatively experience acting on the object and reaching the target. In a second experiment, we first tested whether four other subjects could learn branch leaning after having observed a group-mate performing the task. Despite repeated opportunities to observe the demonstrator, they did not learn to use the pole as a tool. Hence we exposed the latter subjects to individual learning trials and they succeeded in the task. Tool use was not transmitted in the experimental situation, which contrasts with observations in the park. We can conclude that the subjects were not able to recognize the target as such. It is possible that they recognized it and learned the task individually when we alternated the opportunity to act upon the object and to reach the reward. This suggests that these macaques could then have associated the action they exercised upon the pole and the use of the pole as a means to reach the reward.
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Nguyen, N. H., Klein, E. D., & Zentall, T. R. (2005). Imitation of a two-action sequence by pigeons. Psychon Bull Rev, 12(3), 514–518.
Abstract: Developmental psychologists have described imitation as a process that suggests perspective-taking abilities. However, imitative behavior has been found in animals, which are generally not considered capable of taking the perspective of another. Previous studies with birds have demonstrated the imitation of a single response (sometimes referred to as action-level imitation). In the present experiment, we examined the extent to which pigeons would imitate an unfamiliar sequence of two behaviors (sometimes referred to as program-level imitation). Our results indicate that, although there are individual differences, pigeons show a significant tendency to match a demonstrated sequence of behavior involving, first, a response to a treadle (pecking at it or stepping on it) and, second, pushing aside a screen that blocks access to food (a left-vs.-right push).
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Akins, C. K., Klein, E. D., & Zentall, T. R. (2002). Imitative learning in Japanese quail (Coturnix japonica) using the bidirectional control procedure. Anim Learn Behav, 30(3), 275–281.
Abstract: In the bidirectional control procedure, observers are exposed to a conspecific demonstrator responding to a manipulandum in one of two directions (e.g., left vs. right). This procedure controls for socially mediated effects (the mere presence of a conspecific) and stimulus enhancement (attention drawn to a manipulandum by its movement), and it has the added advantage of being symmetrical (the two different responses are similar in topography). Imitative learning is demonstrated when the observers make the response in the direction that they observed it being made. Recently, however, it has been suggested that when such evidence is found with a predominantly olfactory animal, such as the rat, it may result artifactually from odor cues left on one side of the manipulandum by the demonstrator. In the present experiment, we found that Japanese quail, for which odor cues are not likely to play a role, also showed significant correspondence between the direction in which the demonstrator and the observer push a screen to gain access to reward. Furthermore, control quail that observed the screen move, when the movement of the screen was not produced by the demonstrator, did not show similar correspondence between the direction of screen movement observed and that performed by the observer. Thus, with the appropriate control, the bidirectional procedure appears to be useful for studying imitation in avian species.
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Dorrance, B. R., & Zentall, T. R. (2002). Imitation of conditional discriminations in pigeons (Columba livia). J Comp Psychol, 116(3), 277–285.
Abstract: In the present experiments, the 2-action method was used to determine whether pigeons could learn to imitate a conditional discrimination. Demonstrator pigeons (Columba livia) stepped on a treadle in the presence of 1 light and pecked at the treadle in the presence of another light. Demonstration did not seem to affect acquisition of the conditional discrimination (Experiment 1) but did facilitate its reversal of the conditional discrimination (Experiments 2 and 3). The results suggest that pigeons are not only able to learn a specific behavior by observing another pigeon, but they can also learn under which circumstances to perform that behavior. The results have implications for proposed mechanisms of imitation in animals.
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