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Beran, M. J. (2004). Long-term retention of the differential values of Arabic numerals by chimpanzees (Pan troglodytes). Anim. Cogn., 7(2), 86–92.
Abstract: As previously reported (Beran and Rumbaugh, 2001), two chimpanzees used a joystick to collect dots, one-at-a-time, on a computer monitor, and then ended a trial when the number of dots collected was equal to the Arabic numeral presented for the trial. Here, the chimpanzees were presented with the task again after an interval of 6 months and then again after an additional interval of 3.25 years. During each interval, the chimpanzees were not presented with the task, and this allowed an assessment of the extent to which both animals retained the values of each Arabic numeral. Despite lower performance at each retention interval compared to the original study, both chimpanzees performed above chance levels in collecting a quantity of dots equal to the target numeral, one chimpanzee for the numerals 1-7, and the second chimpanzee for the numerals 1-6. For the 3.25-year retention, errors were more dispersed around each target numeral than in the original study, but the chimpanzees' performances again appeared to be based on a continuous representation of magnitude rather than a discrete representation of number. These data provide an experimental demonstration of long-term retention of the differential values of Arabic numerals by chimpanzees.
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Beran, M. J., Pate, J. L., Washburn, D. A., & Rumbaugh, D. M. (2004). Sequential responding and planning in chimpanzees (Pan troglodytes) and rhesus macaques (Macaca mulatta). J Exp Psychol Anim Behav Process, 30(3), 203–212.
Abstract: Chimpanzees (Pan troglodytes) and rhesus macaques (Macaca mulatta) selected either Arabic numerals or colored squares on a computer monitor in a learned sequence. On shift trials, the locations of 2 stimuli were interchanged at some point. More errors were made when this interchange occurred for the next 2 stimuli to be selected than when the interchange was for stimuli later in the sequence. On mask trials, all remaining stimuli were occluded after the 1st selection. Performance exceeded chance levels for only 1 selection after these masks were applied. There was no difference in performance for either stimulus type (numerals or colors). The data indicated that the animals planned only the next selection during these computerized tasks as opposed to planning the entire response sequence.
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Brannon, E. M., & Terrace, H. S. (1998). Ordering of the numerosities 1 to 9 by monkeys. Science, 282(5389), 746–749.
Abstract: A fundamental question in cognitive science is whether animals can represent numerosity (a property of a stimulus that is defined by the number of discriminable elements it contains) and use numerical representations computationally. Here, it was shown that rhesus monkeys represent the numerosity of visual stimuli and detect their ordinal disparity. Two monkeys were first trained to respond to exemplars of the numerosities 1 to 4 in an ascending numerical order (1 --> 2 --> 3 --> 4). As a control for non-numerical cues, exemplars were varied with respect to size, shape, and color. The monkeys were later tested, without reward, on their ability to order stimulus pairs composed of the novel numerosities 5 to 9. Both monkeys responded in an ascending order to the novel numerosities. These results show that rhesus monkeys represent the numerosities 1 to 9 on an ordinal scale.
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Brannon, E. M., & Terrace, H. S. (2000). Representation of the numerosities 1-9 by rhesus macaques (Macaca mulatta). J Exp Psychol Anim Behav Process, 26(1), 31–49.
Abstract: Three rhesus monkeys (Macaca mulatta) were trained to respond to exemplars of 1, 2, 3, and 4 in an ascending, descending, or a nonmonotonic numerical order (1-->2-->3-->4, 4-->3-->2--1, 3-->1-->4-->2). The monkeys were then tested on their ability to order pairs of the novel numerosities 5-9. In Experiment 1, all 3 monkeys ordered novel exemplars of the numerosities 1-4 in ascending or descending order. The attempt to train a nonmonotonic order (3-->1-->4-->2) failed. In Experiment 2A, the 2 monkeys who learned the ascending numerical rule ordered pairs of the novel numerosities 5-9 on unreinforced trials. The monkey who learned the descending numerical rule failed to extrapolate the descending rule to new numerosities. In Experiment 2B all 3 monkeys ordered novel exemplars of pairs of the numerosities 5-9. Accuracy and latency of responding revealed distance and magnitude effects analogous to previous findings with human participants (R. S. Moyer & T. K. Landaeur, 1967). Collectively these studies show that monkeys represent the numerosities 1-9 on at least an ordinal scale.
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Jones, J. E., Antoniadis, E., Shettleworth, S. J., & Kamil, A. C. (2002). A comparative study of geometric rule learning by nutcrackers (Nucifraga columbiana), pigeons (Columba livia), and jackdaws (Corvus monedula). J Comp Psychol, 116(4), 350–356.
Abstract: Three avian species, a seed-caching corvid (Clark's nutcrackers; Nucifraga columbiana), a non-seed-caching corvid (jackdaws; Corvus monedula), and a non-seed-caching columbid (pigeons; Columba livia), were tested for ability to learn to find a goal halfway between 2 landmarks when distance between the landmarks varied during training. All 3 species learned, but jackdaws took much longer than either pigeons or nutcrackers. The nutcrackers searched more accurately than either pigeons or jackdaws. Both nutcrackers and pigeons showed good transfer to novel landmark arrays in which interlandmark distances were novel, but inconclusive results were obtained from jackdaws. Species differences in this spatial task appear quantitative rather than qualitative and are associated with differences in natural history rather than phylogeny.
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Pepperberg, I. M. (2006). Grey parrot numerical competence: a review. Anim. Cogn., 9(4), 377–391.
Abstract: The extent to which humans and nonhumans share numerical competency is a matter of debate. Some researchers argue that nonhumans, lacking human language, possess only a simple understanding of small quantities, generally less than four. Animals that have, however, received some training in human communication systems might demonstrate abilities intermediate between those of untrained nonhumans and humans. Here I review data for a Grey parrot (Psittacus erithacus) that has been shown to quantify sets of up to and including six items (including heterogeneous subsets) using vocal English labels, to comprehend these labels fully, and to have a zero-like concept. Recent research demonstrates that he can also sum small quantities. His success shows that he understands number symbols as abstract representations of real-world collections, and that his sense of number compares favorably to that of chimpanzees and young human children.
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Rumbaugh, D. M., Savage-Rumbaugh, S., & Hegel, M. T. (1987). Summation in the chimpanzee (Pan troglodytes). J Exp Psychol Anim Behav Process, 13(2), 107–115.
Abstract: In this research, we asked whether 2 chimpanzee (Pan troglodytes) subjects could reliably sum across pairs of quantities to select the greater total. Subjects were allowed to choose between two trays of chocolates. Each tray contained two food wells. To select the tray containing the greater number of chocolates, it was necessary to sum the contents of the food wells on each tray. In experiments where food wells contained from zero to four chocolates, the chimpanzees chose the greater value of the summed wells on more than 90% of the trials. In the final experiment, the maximum number of chocolates assigned to a food well was increased to five. Choice of the tray containing the greater sum still remained above 90%. In all experiments, subjects reliably chose the greater sum, even though on many trials a food well on the “incorrect” tray held more chocolates than either single well on the “correct” tray. It was concluded that without any known ability to count, these chimpanzees used some process of summation to combine spatially separated quantities. Speculation regarding the basis for summation includes consideration of perceptual fusion of pairs of quantities and subitization.
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West, R. E., & Young, R. J. (2002). Do domestic dogs show any evidence of being able to count? Anim. Cogn., 5(3), 183–186.
Abstract: Numerical competence has been demonstrated in a wide range of animal species. The level of numerical abilities shown ranges from simple relative numerousness judgements to true counting. In this study we used the preferential looking technique to test whether 11 pet dogs could count. The dogs were presented with three simple calculations: “1+1=2”; “1+1=1”; and “1+1=3”. These calculations were performed by presenting the dogs with treats that were placed behind a screen that allowed manipulation of the outcome of the calculation. When the dogs expected the outcome they spent the same amount of time looking at the result of the calculation as they did on the initial presentation. However, when the result was unexpected dogs spent significantly longer looking at the outcome of the calculation. The results suggest that the dogs were anticipating the outcome of the calculations they observed, thus suggesting that dogs may have a rudimentary ability to count.
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