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Santos, L. R., Barnes, J. L., & Mahajan, N. (2005). Expectations about numerical events in four lemur species (Eulemur fulvus, Eulemur mongoz, Lemur catta and Varecia rubra). Anim. Cogn., 8(4), 253–262.
Abstract: Although much is known about how some primates--in particular, monkeys and apes--represent and enumerate different numbers of objects, very little is known about the numerical abilities of prosimian primates. Here, we explore how four lemur species (Eulemur fulvus, E. mongoz, Lemur catta, and Varecia rubra) represent small numbers of objects. Specifically, we presented lemurs with three expectancy violation looking time experiments aimed at exploring their expectations about a simple 1+1 addition event. In these experiments, we presented subjects with displays in which two lemons were sequentially added behind an occluder and then measured subjects' duration of looking to expected and unexpected outcomes. In experiment 1, subjects looked reliably longer at an unexpected outcome of only one object than at an expected outcome of two objects. Similarly, subjects in experiment 2 looked reliably longer at an unexpected outcome of three objects than at an expected outcome of two objects. In experiment 3, subjects looked reliably longer at an unexpected outcome of one object twice the size of the original than at an expected outcome of two objects of the original size. These results suggest that some prosimian primates understand the outcome of simple arithmetic operations. These results are discussed in light of similar findings in human infants and other adult primates.
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Santos, L. R., & Hauser, M. D. (1999). How monkeys see the eyes: cotton-top tamarins' reaction to changes in visual attention and action. Anim. Cogn., 2(3), 131–139.
Abstract: Among social species, the capacity to detect where another individual is looking is adaptive because gaze direction often predicts what an individual is attending to, and thus what its future actions are likely to be. We used an expectancy violation procedure to determine whether cotton-top tamarins (Saguinus oedipus oedipus) use the direction of another individual's gaze to predict future actions. Subjects were familiarized with a sequence in which a human actor turned her attention toward one of two objects sitting on a table and then reached for that object. Following familiarization, subjects saw two test events. In one test event, the actor gazed at the new object and then reached for that object. From a human perspective, this event is considered consistent with the causal relationship between visual attention and subsequent action, that is, grabbing the object attended to. In the second test event, the actor gazed at the old object, but reached for the new object. This event is considered a violation of expectation. When the actor oriented with both her head-and-eyes, subjects looked significantly longer at the second test event in which the actor reached for the object to which she had not previously oriented. However, there was no difference in looking time between test events when the actor used only her eyes to orient. These findings suggest that tamarins are able to use some combination of head orientation and gaze direction, but not gaze direction alone, to predict the actions of a human agent.
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Santos, L. R., Miller, C. T., & Hauser, M. D. (2003). Representing tools: how two non-human primate species distinguish between the functionally relevant and irrelevant features of a tool. Anim. Cogn., 6(4), 269–281.
Abstract: Few studies have examined whether non-human tool-users understand the properties that are relevant for a tool's function. We tested cotton-top tamarins (Saguinus oedipus) and rhesus macaques (Macaca mulatta) on an expectancy violation procedure designed to assess whether these species make distinctions between the functionally relevant and irrelevant features of a tool. Subjects watched an experimenter use a tool to push a grape down a ramp, and then were presented with different displays in which the features of the original tool (shape, color, orientation) were selectively varied. Results indicated that both species looked longer when a newly shaped stick acted on the grape than when a newly colored stick performed the same action, suggesting that both species perceive shape as a more salient transformation than color. In contrast, tamarins, but not rhesus, attended to changes in the tool's orientation. We propose that some non-human primates begin with a predisposition to attend to a tool's shape and, with sufficient experience, develop a more sophisticated understanding of the features that are functionally relevant to tools.
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Santos, L. R., Pearson, H. M., Spaepen, G. M., Tsao, F., & Hauser, M. D. (2006). Probing the limits of tool competence: experiments with two non-tool-using species (Cercopithecus aethiops and Saguinus oedipus). Anim. Cogn., 9(2), 94–109.
Abstract: Non-human animals vary in their ability to make and use tools. The goal of the present study was to further explore what, if anything, differs between tool-users and non-tool-users, and whether these differences lie in the conceptual or motor domain. We tested two species that typically do not use tools-cotton top tamarins (Saguinus oedipus) and vervet monkeys (Cercopithecus aethiops)-on problems that mirrored those designed for prolific tool users such as chimpanzees. We trained subjects on a task in which they could choose one of two canes to obtain an out-of-reach food reward. After training, subjects received several variations on the original task, each designed to examine a specific conceptual aspect of the pulling problem previously studied in other tool-using species. Both species recognized that effective pulling tools must be made of rigid materials. Subsequent conditions revealed significant species differences, with vervets outperforming tamarins across many conditions. Vervets, but not tamarins, had some recognition of the relationship between a tool's orientation and the position of the food reward, the relationship between a tool's trajectory and the substance that it moves on, and that tools must be connected in order to work properly. These results provide further evidence that tool-use may derive from domain-general, rather than domain-specific cognitive capacities that evolved for tool use per se.
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Santos, L. R., Rosati, A., Sproul, C., Spaulding, B., & Hauser, M. D. (2005). Means-means-end tool choice in cotton-top tamarins (Saguinus oedipus): finding the limits on primates' knowledge of tools. Anim. Cogn., 8(4), 236–246.
Abstract: Most studies of animal tool use require subjects to use one object to gain access to a food reward. In many real world situations, however, animals perform more than one action in sequence to achieve their goals. Of theoretical interest is whether animals have the cognitive capacity to recognize the relationship between consecutive action sequences in which there may be one overall goal and several subgoals. Here we ask if cotton-top tamarins, a species that in captivity uses tools to solve means-end problems, can go one step further and use a sequence of tools (means) to obtain food (end). We first trained subjects to use a pulling tool to obtain a food reward. After this initial training, subjects were presented with problems in which one tool had to be used in combination with a second in order to obtain food. Subjects showed great difficulty when two tools were required to obtain the food reward. Although subjects attended to the connection between the tool and food reward, they ignored the physical connection between the two tools. After training on a two-tool problem, we presented subjects with a series of transfer tests to explore if they would generalize to new types of connections between the tools. Subjects readily transferred to new connections. Our results therefore provide the first evidence to date that tamarins can learn to solve problems involving two tools, but that they do so only with sufficient training.
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