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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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Beran, M. J. (2007). Rhesus monkeys (Macaca mulatta) succeed on a computerized test designed to assess conservation of discrete quantity. Anim. Cogn., 10(1), 37–45.
Abstract: Conservation of quantity occurs through recognition that changes in the physical arrangement of a set of items do not change the quantity of items in that set. Rhesus monkeys (Macaca mulatta) were presented with a computerized quantity judgment task. Monkeys were rewarded for selecting the greater quantity of items in one of two horizontal arrays of items on the screen. On some trials, after a correct selection, no reward was given but one of the arrays was manipulated. In some cases, this manipulation involved moving items closer together or farther apart to change the physical arrangement of the array without changing the quantity of items in the array. In other cases, additional items were added to the initially smaller array so that it became quantitatively larger. Monkeys then made another selection from the two rows of items. Monkeys were sensitive to these manipulations, changing their selections when the number of items in the rows changed but not when the arrangement only was changed. Therefore, monkeys responded on the basis of the quantity of items, and they were not distracted by non-quantitative manipulations of the sets.
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Jordan, K. E., & Brannon, E. M. (2006). Weber's Law influences numerical representations in rhesus macaques (Macaca mulatta). Anim. Cogn., 9(3), 159–172.
Abstract: We present the results of two experiments that probe the ability of rhesus macaques to match visual arrays based on number. Three monkeys were first trained on a delayed match-to-sample paradigm (DMTS) to match stimuli on the basis of number and ignore continuous dimensions such as element size, cumulative surface area, and density. Monkeys were then tested in a numerical bisection experiment that required them to indicate whether a sample numerosity was closer to a small or large anchor value. Results indicated that, for two sets of anchor values with the same ratio, the probability of choosing the larger anchor value systematically increased with the sample number and the psychometric functions superimposed. A second experiment employed a numerical DMTS task in which the choice values contained an exact numerical match to the sample and a distracter that varied in number. Both accuracy and reaction time were modulated by the ratio between the correct numerical match and the distracter, as predicted by Weber's Law.
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Lewis, K. P., Jaffe, S., & Brannon, E. M. (2005). Analog number representations in mongoose lemurs (Eulemur mongoz): evidence from a search task. Anim. Cogn., 8(4), 247–252.
Abstract: A wealth of data demonstrating that monkeys and apes represent number have been interpreted as suggesting that sensitivity to number emerged early in primate evolution, if not before. Here we examine the numerical capacities of the mongoose lemur (Eulemur mongoz), a member of the prosimian suborder of primates that split from the common ancestor of monkeys, apes and humans approximately 47-54 million years ago. Subjects observed as an experimenter sequentially placed grapes into an opaque bucket. On half of the trials the experimenter placed a subset of the grapes into a false bottom such that they were inaccessible to the lemur. The critical question was whether lemurs would spend more time searching the bucket when food should have remained in the bucket, compared to when they had retrieved all of the food. We found that the amount of time lemurs spent searching was indicative of whether grapes should have remained in the bucket, and furthermore that lemur search time reliably differentiated numerosities that differed by a 1:2 ratio, but not those that differed by a 2:3 or 3:4 ratio. Finally, two control conditions determined that lemurs represented the number of food items, and neither the odor of the grapes, nor the amount of grape (e.g., area) in the bucket. These results suggest that mongoose lemurs have numerical representations that are modulated by Weber's Law.
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Harris, E. H., & Washburn, D. A. (2005). Macaques' (Macaca mulatta) use of numerical cues in maze trials. Anim. Cogn., 8(3), 190–199.
Abstract: We tested the ability of number-trained rhesus monkeys to use Arabic numeral cues to discriminate between different series of maze trials and anticipate the final trial in each series. The monkeys' prior experience with numerals also allowed us to investigate spontaneous transfer between series. A total of four monkeys were tested in two experiments. In both experiments, the monkeys were trained on a computerized task consisting of three reinforced maze trials followed by one nonreinforced trial. The goal of the maze was an Arabic numeral 3, which corresponded to the number of reinforced maze trials in the series. In experiment 1 (n=2), the monkeys were given probe trials of the numerals 2 and 4 and in experiment 2 (n=2), they were given probe trials of the numerals 2-8. The monkeys receiving the probe trials 2 and 4 showed some generalization to the new numerals and developed a pattern of performing more slowly on the nonreinforced trial than the reinforced trial before it for most series, indicating the use of the changing numeral cues to anticipate the nonreinforced trial. The monkeys receiving probe trials of the numerals 2-8 did not predict precisely when the nonreinforced trial would occur in each series, but they did incorporate the changing numerals into their strategy for performing the task. This study provides the first evidence that number-trained monkeys can use Arabic numerals to perform a task involving sequential presentations.
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Ferkin, M. H., Pierce, A. A., Sealand, R. O., & Delbarco-Trillo, J. (2005). Meadow voles, Microtus pennsylvanicus, can distinguish more over-marks from fewer over-marks. Anim. Cogn., 8(3), 182–189.
Abstract: Is it possible that voles have a sense of number? To address this question, we determined whether voles discriminate between two different scent-marking individuals and identify the individual whose scent marks was on top more often than the other individual. We tested whether voles show a preference for the individual whose scent marks was on top most often. If so, the simplest explanation was that voles can make a relative size judgement-such as distinguishing an area containing more of one individual's over-marks as compared to less of another individual's over-marks. We found that voles respond preferentially to the donor that provided a greater number of over-marks as compared to the donor that provided a lesser number of over-marks. Thus, we concluded that voles might display the capacity for relative numerousness. Interestingly, female voles were better able than male voles to distinguish between small differences in the relative number of over-marks by the two scent donors.
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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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Uller, C., Jaeger, R., Guidry, G., & Martin, C. (2003). Salamanders ( Plethodon cinereus) go for more: rudiments of number in an amphibian. Anim. Cogn., 6(2), 105–112.
Abstract: Techniques traditionally used in developmental research with infants have been widely used with nonhuman primates in the investigation of comparative cognitive abilities. Recently, researchers have shown that human infants and monkeys select the larger of two numerosities in a spontaneous forced-choice discrimination task. Here we adopt the same method to assess in a series of experiments spontaneous choice of the larger of two numerosities in a species of amphibian, red-backed salamanders ( Plethodon cinereus). The findings indicate that salamanders “go for more,” just like human babies and monkeys. This rudimentary capacity is a type of numerical discrimination that is spontaneously present in this amphibian.
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Kiltie, R. A., Fan, J., & Laine, A. F. (1995). A wavelet-based metric for visual texture discrimination with applications in evolutionary ecology. Math Biosci, 126(1), 21–39.
Abstract: Much work on natural and sexual selection is concerned with the conspicuousness of visual patterns (textures) on animal and plant surfaces. Previous attempts by evolutionary biologists to quantify apparency of such textures have involved subjective estimates of conspicuousness or statistical analyses based on transect samples. We present a method based on wavelet analysis that avoids subjectivity and that uses more of the information in image textures than transects do. Like the human visual system for texture discrimination, and probably like that of other vertebrates, this method is based on localized analysis of orientation and frequency components of the patterns composing visual textures. As examples of the metric's utility, we present analyses of crypsis for tigers, zebras, and peppered moth morphs.
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