Catherine F. Talbot

Putting the elephant back in the herd: elephant relative quantity judgments match those of other species

The ability to discriminate between quantities has been observed in many species. Typically, when an animal is given a choice between two sets of food, accurate performance (i.e., choosing the larger amount) decreases as the ratio between two quantities increases. A recent study reported that elephants did not exhibit ratio effects, suggesting that elephants may process quantitative information in a qualitatively different way from all other nonhuman species that have been tested (Irie-Sugimoto et al. in Anim Cogn 12:193–199, 2009). However, the results of this study were confounded by several methodological issues. We tested two African elephants (Loxodonta africana) to more thoroughly investigate relative quantity judgment in this species. In contrast to the previous study, we found evidence of ratio effects for visible and nonvisible sequentially presented sets of food. Thus, elephants appear to represent and compare quantities in much the same way as other species, including humans when they are prevented from counting. Performance supports an accumulator model in which quantities are represented as analog magnitudes. Furthermore, we found no effect of absolute magnitude on performance, providing support against an object-file model explanation of quantity judgment.

Squirrel monkeys' response to inequitable outcomes indicates a behavioural convergence within the primates

Although several primates respond negatively to inequity, it is unknown whether this results from homology or convergent processes. Behaviours shared within a taxonomic group are often assumed to be homologous, yet this distinction is important for a better understanding of the function of the behaviour. Previous hypotheses have linked cooperation and inequity responses. Supporting this, all species in which inequity responses have been documented are cooperative. In this study, we tested this hypothesis by investigating the response to inequity in squirrel monkeys, which share a phylogenetic family with capuchin monkeys, but do not cooperate extensively. Subjects exchanged tokens to receive food rewards in conditions in which the level of effort required and reward received varied. Squirrel monkeys did not respond negatively to inequity. However, the monkeys were sensitive to the variation present in the task; male subjects showed a contrast effect and, as in previous studies, subjects were more sensitive to differences in reward in the context of a task than when rewards were given for free. Taken with other results, these results support the hypothesis that a negative response to inequity evolved convergently in primates, probably as a mechanism for evaluating outcomes relative to ones partners in cooperative species.

Responses to the Assurance game in monkeys, apes, and humans using equivalent procedures

There is great interest in the evolution of economic behavior. In typical studies, species are asked to play one of a series of economic games, derived from game theory, and their responses are compared. The advantage of this approach is the relative level of consistency and control that emerges from the games themselves; however, in the typical experiment, procedures and conditions differ widely, particularly between humans and other species. Thus, in the current study, we investigated how three primate species, capuchin monkeys, chimpanzees, and humans, played the Assurance (or Stag Hunt) game using procedures that were, to the best of our ability, the same across species, particularly with respect to training and pretesting. Our goal was to determine what, if any, differences existed in the ways in which these species made decisions in this game. We hypothesized differences along phylogenetic lines, which we found. However, the species were more similar than might be expected. In particular, humans who played using “nonhuman primate-friendly” rules did not behave as is typical. Thus, we find evidence for similarity in decision-making processes across the order Primates. These results indicate that such comparative studies are possible and, moreover, that in any comparison rating species’ relative abilities, extreme care must be taken in ensuring that one species does not have an advantage over the others due to methodological procedures.

Orangutans (Pongo pygmaeus) Do Not Form Expectations Based on Their Partner’s Outcomes

Several primate species form expectations based on others’ outcomes, responding negatively when their outcomes differ from their partners’. The function and evolutionary pathway of this behavior are unknown, in part because all of the species which have been tested thus far share traits related to a gregarious lifestyle, intelligence, and cooperativeness. Our goal was to test whether inequity is a homology among primates or a convergence by comparing one species known to show social comparisons, the chimpanzee, to another great ape which differs on several of these life history characteristics. Using a protocol identical to one used previously with chimpanzees, we tested whether orangutans, an intelligent but predominantly solitary species with few opportunities to cooperate, responded similarly. To allow for a strong comparison with chimpanzees (and other species), we used socially housed adults of both sexes, tested with members of their social group. Orangutans did not respond negatively to inequity, supporting previous findings and indicating that inequity responses in apes are likely a convergence based on either sociality or cooperative tendency. These results in such closely related species highlight the need for additional comparative studies to understand better the function and evolution of social behaviors.

Different Responses to Reward Comparisons by Three Primate Species

Background Recently, much attention has been paid to the role of cooperative breeding in the evolution of behavior. In many measures, cooperative breeders are more prosocial than non-cooperatively breeding species, including being more likely to actively share food. This is hypothesized to be due to selective pressures specific to the interdependency characteristic of cooperatively breeding species. Given the high costs of finding a new mate, it has been proposed that cooperative breeders, unlike primates that cooperate in other contexts, should not respond negatively to unequal outcomes between themselves and their partner. However, in this context such pressures may extend beyond cooperative breeders to other species with pair-bonding and bi-parental care. Methods Here we test the response of two New World primate species with different parental strategies to unequal outcomes in both individual and social contrast conditions. One species tested was a cooperative breeder (Callithrix spp.) and the second practiced bi-parental care (Aotus spp.). Additionally, to verify our procedure, we tested a third confamilial species that shows no such interdependence but does respond to individual (but not social) contrast (Saimiri spp.). We tested all three genera using an established inequity paradigm in which individuals in a pair took turns to gain rewards that sometimes differed from those of their partners. Conclusions None of the three species tested responded negatively to inequitable outcomes in this experimental context. Importantly, the Saimiri spp responded to individual contrast, as in earlier studies, validating our procedure. When these data are considered in relation to previous studies investigating responses to inequity in primates, they indicate that one aspect of cooperative breeding, pair-bonding or bi-parental care, may influence the evolution of these behaviors. These results emphasize the need to study a variety of species to gain insight in to how decision-making may vary across social structures.

Inequity Responses in Nonhuman Animals

Aversion to inequity is a universal human trait, but recent evidence in other primate and nonprimate species suggests that humans are not the only species to respond negatively to unequal outcomes. Although only about a dozen species have been tested thus far, the emerging phylogenetic patterns help us to understand the evolution of responses to inequity. Negative responses to inequity are not universal, even among the primates, nor are they based upon social tolerance, social group size, or brain size. Instead, what we see is that all of the species that respond to inequity habitually cooperate with nonkin in various contexts, including food sharing, cooperative hunting, coalitions, and alliances. This indicates that inequity responses emerged in conjunction with cooperation. Their likely function is to compare one’s own effort and payoff with that of a partner, and therefore help individuals determine the value of a cooperative partner. In other words, negative reactions to inequity may stabilize cooperation by helping individuals recognize when a cooperative partner is getting more than their “fair share,” which is a cue that it may be time to find a new, more equitable, partner. These negative responses to inequity do not necessarily involve rational or even conscious decision-making, suggesting that they are widespread among animals that cooperate. We end by speculating what these data mean for our understanding of the evolution of fairness in humans, and what this says about the human sense of morality.

The Influence of Reward Quality and Quantity and Spatial Proximity on the Responses to Inequity and Contrast in Capuchin Monkeys (Cebus [Sapajus] apella).

Recent evidence within the field of comparative psychology has demonstrated that small differences in procedure may lead to significant differences in outcome. Therefore, failing to fully explore the impact of different contexts on a behavior limits our ability to fully understand that behavior. A behavior that has exhibited substantial variation, both within and across studies, is animals’ responses to violations of their expectations, either when expectations were based on another’s outcome (inequity) or one’s previous outcome (contrast). We explored this further in capuchin monkeys, focusing on the following 2 factors that often vary in such tests but have not yet been rigorously explored: the relative values of the food rewards and the degree of separation of the subjects. Concerning the first, we examined responses to violation of expectations when the difference between what was expected (or what the partner got) and what was received differed in either quality or quantity. Concerning the second, we compared responses when the 2 individuals were separated by a clear partition (barrier condition) versus sharing the same enclosure without the partition (no-barrier condition). Our results suggest that responses to inequity are most likely to emerge when the food received is low-value food, regardless of the difference between the actual and the expected outcome. However, capuchins did not respond differently to different quantities of rewards, nor did the degree of separation between subjects significantly affect results. We consider the implications of this work for both studies of violation of expectation and other cognitive and behavioral tasks.