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Hare, B., Addessi, E., Call, J., Tomasello, M., & Visalberghi, E. (2003). Do capuchin monkeys, Cebus apella, know what conspecifics do and do not see? Anim. Behav., 65(1), 131–142.
Abstract: Capuchin monkeys were tested in five experiments in which two individuals competed over food. When given a choice between retrieving a piece of food that was visible or hidden from the dominant, subordinate animals preferred to retrieve hidden food. This preference is consistent with the hypotheses that either (1) the subordinate knew what the dominant could and could not see or (2) the subordinate was monitoring the behaviour of the dominant and avoiding the piece of food that it approached. To test between these alternatives, we released subordinates with a slight head start forcing them to make their choice (between a piece of food hidden or visible to the dominant) before the dominant entered the area. Unlike chimpanzees, Pan troglodytes, subordinates that were given a head start did not preferentially approach hidden pieces of food first. Therefore, our experiments provide little support for the hypothesis that capuchin monkeys are sensitive to what another individual does or does not see. We compare our results with those obtained with chimpanzees in the same paradigm and discuss the evolution of primate social cognition. Copyright 2003 The Association for the Study of Animal Behaviour. Published by Elsevier Science Ltd. All rights reserved.
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Hare, B., Call, J., Agnetta, B., & Tomasello, M. (2000). Chimpanzees know what conspecifics do and do not see. Anim. Behav., 59(4), 771–785.
Abstract: We report a series of experiments on social problem solving in chimpanzees, Pan troglodytes. In each experiment a subordinate and a dominant individual were put into competition over two pieces of food. In all experiments dominants obtained virtually all of the foods to which they had good visual and physical access. However, subordinates were successful quite often in three situations in which they had better visual access to the food than the dominant, for example, when the food was positioned so that only the subordinate (and not the dominant) could see it. In some cases, the subordinate might have been monitoring the behaviour of the dominant directly and simply avoided the food that the dominant was moving towards (which just happened to be the one it could see). In other cases, however, we ruled out this possibility by giving subordinates a small headstart and forcing them to make their choice (to go to the food that both competitors could see, or the food that only they could see) before the dominant was released into the area. Together with other recent studies, the present investigation suggests that chimpanzees know what conspecifics can and cannot see, and, furthermore, that they use this knowledge to devise effective social-cognitive strategies in naturally occurring food competition situations.
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Hare, B., Call, J., & Tomasello, M. (2001). Do chimpanzees know what conspecifics know? Anim. Behav., 61(1), 139–151.
Abstract: We conducted three experiments on social problem solving by chimpanzees, Pan troglodytes. In each experiment a subordinate and a dominant individual competed for food, which was placed in various ways on the subordinate's side of two opaque barriers. In some conditions dominants had not seen the food hidden, or food they had seen hidden was moved elsewhere when they were not watching (whereas in control conditions they saw the food being hidden or moved). At the same time, subordinates always saw the entire baiting procedure and could monitor the visual access of their dominant competitor as well. If subordinates were sensitive to what dominants did or did not see during baiting, they should have preferentially approached and retrieved the food that dominants had not seen hidden or moved. This is what they did in experiment 1 when dominants were either uninformed or misinformed about the food's location. In experiment 2 subordinates recognized, and adjusted their behaviour accordingly, when the dominant individual who witnessed the hiding was replaced with another dominant individual who had not witnessed it, thus demonstrating their ability to keep track of precisely who has witnessed what. In experiment 3 subordinates did not choose consistently between two pieces of hidden food, one of which dominants had seen hidden and one of which they had not seen hidden. However, their failure in this experiment was likely to be due to the changed nature of the competition under these circumstances and not to a failure of social-cognitive skills. These findings suggest that at least in some situations (i.e. competition with conspecifics) chimpanzees know what conspecifics have and have not seen (do and do not know), and that they use this information to devise effective social-cognitive strategies. Copyright 2001 The Association for the Study of Animal Behaviour.
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Byrnl, R. W., & Tomasello, M. (1995). Do rats ape? Anim. Behav., 50(5), 1417–1420.
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Tomasello, M., Call, J., & Hare, B. (1998). Five primate species follow the visual gaze of conspecifics. Anim. Behav., 55(4), 1063–1069.
Abstract: Individuals from five primate species were tested experimentally for their ability to follow the visual gaze of conspecifics to an outside object. Subjects were from captive social groups of chimpanzees,Pan troglodytes, sooty mangabeys,Cercocebus atys torquatus, rhesus macaques,Macaca mulatta, stumptail macaques,M. arctoides, and pigtail macaques,M. nemestrina. Experimental trials consisted of an experimenter inducing one individual to look at food being displayed, and then observing the reaction of another individual (the subject) that was looking at that individual (not the food). Control trials consisted of an experimenter displaying the food in an identical manner when the subject was alone. Individuals from all species reliably followed the gaze of conspecifics, looking to the food about 80% of the time in experimental trials, compared with about 20% of the time in control trials. Results are discussed in terms of both the proximate mechanisms that might be involved and the adaptive functions that might be served by gaze-following.
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Tomasello, M., Hare, B., & Fogleman, T. (2001). The ontogeny of gaze following in chimpanzees, Pan troglodytes, and rhesus macaques, Macaca mulatta. Anim. Behav., 61(2), 335–343.
Abstract: Primates follow the gaze direction of conspecifics to outside objects. We followed the ontogeny of this social-cognitive skill for two species: rhesus macaques and chimpanzees. In the first two experiments, using both a cross-sectional and a longitudinal design, we exposed individuals of different ages to a human looking in a specified direction. Rhesus infants first began reliably to follow the direction of this gaze at the end of the early infancy period, at about 5.5 months of age. Chimpanzees did not reliably follow human gaze until 3-4 years; this corresponds to the latter part of the late infancy period for this species. In the third experiment we exposed individuals of the same two species to a human repeatedly looking to the same location (with no special object at that location) to see if subjects would learn to ignore the looks. Only adults of the two species diminished their gaze-following behaviour over trials. This suggests that in the period between infancy and adulthood individuals of both species come to integrate their gaze-following skills with their more general social-cognitive knowledge about other animate beings and their behaviour, and so become able to deploy their gaze-following skills in a more flexible manner.
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Herrmann, E., Melis, A. P., & Tomasello, M. (2006). Apes' use of iconic cues in the object-choice task. Anim. Cogn., 9(2), 118–130.
Abstract: In previous studies great apes have shown little ability to locate hidden food using a physical marker placed by a human directly on the target location. In this study, we hypothesized that the perceptual similarity between an iconic cue and the hidden reward (baited container) would help apes to infer the location of the food. In the first two experiments, we found that if an iconic cue is given in addition to a spatial/indexical cue – e.g., picture or replica of a banana placed on the target location – apes (chimpanzees, bonobos, orangutans, gorillas) as a group performed above chance. However, we also found in two further experiments that when iconic cues were given on their own without spatial/indexical information (iconic cue held up by human with no diagnostic spatial/indexical information), the apes were back to chance performance. Our overall conclusion is that although iconic information helps apes in the process of searching hidden food, the poor performance found in the last two experiments is due to apes' lack of understanding of the informative (cooperative) communicative intention of the experimenter.
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Buttelmann, D., Call, J., & Tomasello, M. (2007). Behavioral cues that great apes use to forage for hidden food. Anim. Cogn., .
Abstract: We conducted three studies to examine whether the four great ape species (chimpanzees, bonobos, gorillas, and orangutans) are able to use behavioral experimenter-given cues in an object-choice task. In the subsequent experimental conditions subjects were presented with two eggs, one of which contained food and the other did not. In Study 1 the experimenter examined both eggs by smelling or shaking them, but only made a failed attempt to open (via biting) the egg containing food. In a control condition, the experimenter examined and attempted to open both eggs, but in reverse order to control for stimulus enhancement. The apes significantly preferred the egg that was first examined and then bitten, but had no preference in a baseline condition in which there were no cues. In Study 2, we investigated whether the apes could extend this ability to cues not observed in apes so far (i.e., attempting to pull apart the egg), as well as whether they made this discrimination based on the function of the action the experimenter performed. Subjects significantly preferred eggs presented with this novel cue, but did not prefer eggs presented with a novel but functionally irrelevant action. In Study 3, apes did not interpret human actions as cues to food-location when they already knew that the eggs were empty. Thus, great apes were able to use a variety of experimenter-given cues associated with foraging actions to locate hidden food and thereby were partially sensitive to the general purpose underlying these actions.
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Riedel, J., Buttelmann, D., Call, J., & Tomasello, M. (2006). Domestic dogs (Canis familiaris) use a physical marker to locate hidden food. Anim. Cogn., 9(1), 27–35.
Abstract: Dogs can use the placement of an arbitrary marker to locate hidden food in an object-choice situation. We tested domestic dogs (Canis familiaris) in three studies aimed at pinning down the relative contributions of the human's hand and the marker itself. We baited one of two cups (outside of the dogs' view) and gave the dog a communicative cue to find the food. Study 1 systematically varied dogs' perceptual access to the marker placing event, so that dogs saw either the whole human, the hand only, the marker only, or nothing. Follow-up trials investigated the effect of removing the marker before the dog's choice. Dogs used the marker as a communicative cue even when it had been removed prior to the dog's choice and attached more importance to this cue than to the hand that placed it although the presence of the hand boosted performance when it appeared together with the marker. Study 2 directly contrasted the importance of the hand and the marker and revealed that the effect of the marker diminished if it had been associated with both cups. In contrast touching both cups with the hand had no effect on performance. Study 3 investigated whether the means of marker placement (intentional or accidental) had an effect on dogs' choices. Results showed that dogs did not differentiate intentional and accidental placing of the marker. These results suggest that dogs use the marker as a genuine communicative cue quite independently from the experimenter's actions.
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Call, J., Carpenter, M., & Tomasello, M. (2005). Copying results and copying actions in the process of social learning: chimpanzees (Pan troglodytes) and human children (Homo sapiens). Anim. Cogn., 8(3), 151–163.
Abstract: There is currently much debate about the nature of social learning in chimpanzees. The main question is whether they can copy others' actions, as opposed to reproducing the environmental effects of these actions using their own preexisting behavioral strategies. In the current study, chimpanzees (Pan troglodytes) and human children (Homo sapiens) were shown different demonstrations of how to open a tube-in both cases by a conspecific. In different experimental conditions, demonstrations consisted of (1) action only (the actions necessary to open the tube without actually opening it); (2) end state only (the open tube, without showing any actions); (3) both of these components (in a full demonstration); or (4) neither of these components (in a baseline condition). In the first three conditions subjects saw one of two different ways that the tube could open (break in middle; caps off ends). Subjects' behavior in each condition was assessed for how often they opened the tube, how often they opened it in the same location as the demonstrator, and how often they copied the demonstrator's actions or style of opening the tube. Whereas chimpanzees reproduced mainly the environmental results of the demonstrations (emulation), human children often reproduced the demonstrator's actions (imitation). Because the procedure used was similar in many ways to the procedure that Meltzoff (Dev Psych 31:1, 1995) used to study the understanding of others' unfulfilled intentions, the implications of these findings with regard to chimpanzees' understanding of others' intentions are also discussed.
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