Beran, M. J., Smith, J. D., & Perdue, B. M. (2013). Language-Trained Chimpanzees (Pan troglodytes) Name What They Have Seen but Look First at What They Have Not Seen. Psychol Sci, .
Abstract: Metacognition can be defined as knowing what one knows, and the question of whether nonhuman animals are metacognitive has driven an intense debate. We tested 3 language-trained chimpanzees in an information-seeking task in which the identity of a food item was the critical piece of information needed to obtain the food. The chimpanzees could either report the identity of the food immediately or first check a container in which the food had been hidden. In two experiments, the chimpanzees were significantly more likely to visit the container first on trials in which they could not know its contents but were more likely to just name the food item without looking into the container on trials in which they had seen its contents. Thus, chimpanzees showed efficient information-seeking behavior that suggested they knew what they had or had not already seen when it was time to name a hidden item.
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Beran, M. J., Smith, J. D., Redford, J. S., & Washburn, D. A. (2006). Rhesus macaques (Macaca mulatta) monitor uncertainty during numerosity judgments. J Exp Psychol Anim Behav Process, 32(2), 111–119.
Abstract: Two rhesus macaques (Macaca mulatta) judged arrays of dots on a computer screen as having more or fewer dots than a center value that was never presented in trials. After learning a center value, monkeys were given an uncertainty response that let them decline to make the numerosity judgment on that trial. Across center values (3-7), errors occurred most often for sets adjacent in numerosity to the center value. The monkeys also used the uncertainty response most frequently on these difficult trials. A 2nd experiment showed that monkeys' responses reflected numerical magnitude and not the surface-area illumination of the displays. This research shows that monkeys' uncertainty-monitoring capacity extends to the domain of numerical cognition. It also shows monkeys' use of the purest uncertainty response possible, uncontaminated by any secondary motivator.
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Maynard Smith, J. (1982). Evolution and the Theory of Games.
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Maynard Smith, J., & Price, G. R. (1973). The Logic of Animal Conflict. Nature, 246, 15–18.
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Nuñez, C. M. V., Adelman, J. S., Smith, J., Gesquiere, L. R., & Rubenstein, D. I. (2014). Linking social environment and stress physiology in feral mares (Equus caballus): Group transfers elevate fecal cortisol levels. General and Comparative Endocrinology, 196, 26–33.
Abstract: Abstract Feral horses (Equus caballus) have a complex social structure, the stability of which is important to their overall health. Behavioral and demographic research has shown that decreases in group (or band) stability reduce female fitness, but the potential effects on the physiological stress response have not been demonstrated. To fully understand how band stability affects group-member fitness, we need to understand not only behavioral and demographic, but also physiological consequences of decreases to that stability. We studied group changes in feral mares (an activity that induces instability, including both male and female aggression) on Shackleford Banks, NC. We found that mares in the midst of changing groups exhibit increased fecal cortisol levels. In addition, mares making more group transfers show higher levels of cortisol two weeks post-behavior. These results offer insights into how social instability is integrated into an animal’s physiological phenotype. In addition, our results have important implications for feral horse management. On Shackleford Banks, mares contracepted with porcine zona pellucida (PZP) make approximately 10 times as many group changes as do untreated mares. Such animals may therefore be at higher risk of chronic stress. These results support the growing consensus that links between behavior and physiological stress must be taken into account when managing for healthy, functional populations.
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Pinkus, S., Smith, J., & Jolly, A. (2006). Feeding Competition Between Introduced Eulemur fulvus and Native Lemur catta During the Birth Season at Berenty Reserve, Southern Madagascar. In Ringtailed Lemur Biology (pp. 119–140).
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Smith, J. E., Kolowski, J. M., Graham, K. E., Dawes, S. E., & Holekamp, K. E. (2008). Social and ecological determinants of fission-fusion dynamics in the spotted hyaena. Anim. Behav., 76(3), 619–636.
Abstract: Theory predicts that individuals living in fission-fusion societies, in which group members frequently change subgroups, should modify grouping patterns in response to varying social and environmental conditions. Spotted hyaenas, Crocuta crocuta, are long-lived carnivores that reside in permanent social groups called clans. Clans are complex, fission-fusion societies in which individual members travel, rest and forage in subgroups that frequently change composition. We studied two clans in Kenya to provide the first detailed description of fission-fusion dynamics in this species. Because social and ecological circumstances can influence the cohesiveness of animal societies, we evaluated the extent to which specific circumstances promote the formation of subgroups of various sizes. We found that cooperative defence of shared resources during interclan competition and protection from lions were cohesive forces that promoted formation of large subgroups. We also tested hypotheses suggesting factors limiting subgroup size. Mothers with small cubs avoided conspecifics, thereby reducing infanticide risk. Victims of aggression either reconciled fights or separated from former opponents to reduce the immediate costs of escalated aggression in the absence of food. As predicted by the ecological constraints hypothesis, hyaenas adjusted their grouping patterns over both short and long time scales in response to feeding competition. Crocuta were most gregarious during periods of abundant prey, joined clanmates at ephemeral kills in numbers that correlated with the energetic value of the prey and gained the most energy when foraging alone because cooperative hunting attracted numerous competitors. Overall, our findings indicate that resource limitation constrains grouping in this species.
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Smith, J. E., Van Horn, R. C., Powning, K. S., Cole, A. R., Graham, K. E., Memenis, S. K., et al. (2010). Evolutionary forces favoring intragroup coalitions among spotted hyenas and other animals. Behav. Ecol., 21(2), 284–303.
Abstract: Coalitionary support in agonistic interactions represents cooperation because intervening in a fight is potentially costly to the donor of support but benefits the recipient. Here, we first review the characteristics of, and evolutionary forces favoring, intragroup coalitions in 49 species and find that patterns of intragroup coalition formation are remarkably similar between primates and nonprimates. We then test hypotheses suggesting kin selection, reciprocal altruism, and direct benefits as adaptive explanations for coalitionary interventions among adult female spotted hyenas (Crocuta crocuta) belonging to a large social group in Kenya. As predicted by kin selection theory, females supported close kin most often, and the density (connectedness) of cooperation networks increased with genetic relatedness. Nevertheless, kinship failed to protect females from coalitionary attacks. We found no evidence of enduring alliances based on reciprocal support among unrelated adult females. Instead, donors generally minimized costs to themselves, intervening most often during low-intensity fights and when feeding opportunities were unavailable. Females also gained direct benefits from directing coalitionary attacks toward subordinates. Finally, females monitored the number of dominant bystanders in the “audience” at fights and modified their level of cooperation based on this knowledge. Overall, hyenas made flexible decisions regarding whether or not to intervene in fights, modifying their tendency to cooperate based on multiple types of information about their immediate social and ecological environments. Taken together, these findings indicate that the combined evolutionary forces of kin selection and direct benefits derived from reinforcing the status quo drive coalitionary interventions among adult female spotted hyenas.
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Smith, J. M., & Parker, G. A. (1976). The logic of asymmetric contests. Anim. Behav., 24(1), 159–175.
Abstract: A theoretical analysis is made of the evolution of behavioural strategies in contest situations. It is assumed that behaviour will evolve so as to maximize individual fitness. If so, a population will evolve an [`]evolutionarily stable strategy', or ESS, which can be defined as a strategy such that, if all members of a population adopt it, no [`]mutant' strategy can do better. A number of simple models of contest situations are analysed from this point of view. It is concluded that in [`]symmetric' contests the ESS is likely to be a [`]mixed' strategy; that is, either the population will be genetically polymorphic or individuals will be behaviourally variable. Most real contests are probably asymmetric, either in pay-off to the contestants, or in size or weapons, or in some [`]uncorrelated' fashion; i.e. in a fashion which does not substantially bias either the pay-offs or the likely outcome of an escalated contest. An example of an uncorrelated asymmetry is that between the [`]discoverer' of a resource and a [`]late-comer'. It is shown that the ESS in asymmetric contests will usually be to permit the asymmetric cue to settle the contest without escalation. Escalated contests will, however, occur if information to the contestants about the asymmetry is imperfect.
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Washburn, D. A., Smith, J. D., & Shields, W. E. (2006). Rhesus monkeys (Macaca mulatta) immediately generalize the uncertain response. J Exp Psychol Anim Behav Process, 32(2), 185–189.
Abstract: Rhesus monkeys (Macaca mulatta) have learned, like humans, to use an uncertain response adaptively under test conditions that create uncertainty, suggesting a metacognitive process by which human and nonhuman primates may monitor their confidence and alter their behavior accordingly. In this study, 4 rhesus monkeys generalized their use of the uncertain response, without additional training, to 2 familiar tasks (2-choice discrimination learning and mirror-image matching to sample) that predictably and demonstrably produce uncertainty. The monkeys were significantly less likely to use the uncertain response on trials in which the answer might be known. These results indicate that monkeys, like humans, know when they do not know and that they can learn to use a symbol as a generalized means for indicating their uncertainty.
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