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A key goal of the field of endocrinology has been to understand the hormonal mechanisms that drive social behavior and influence reactions to others, such as oxytocin. However, it has sometimes been challenging to understand which aspects and influences of hormonal action are conserved and common among mammalian species, and which effects differ based on features of these species, such as social system. This challenge has been exacerbated by a focus on a relatively small number of traditional model species. In this review, we first demonstrate the benefits of using non-traditional models for the study of hormones, with a focus on oxytocin as a case study in adding species with diverse social systems. We then expand our discussion to explore differing effects of oxytocin (and its response to behavior) within a species, with a particular focus on relationship context and social environment among primate species. Finally, we suggest key areas for future exploration of oxytocin's action centrally and peripherally, and how non-traditional models can be an important resource for understanding the breadth of oxytocin's potential effects., Competing Interests: The authors declare that the research was conducted in the absence of any commercial or financial relationships that could be construed as a potential conflict of interest. The author(s) declared that they were an editorial board member of Frontiers, at the time of submission. This had no impact on the peer review process and the final decision., (Copyright © 2024 Sosnowski and Brosnan.)

The ability to quickly perceive others' rank minimizes costs by helping individuals behave appropriately when interacting with strangers. Indeed, humans and at least some other species can quickly determine strangers' rank or dominance based only on physical features without observing others' interactions or behavior. Nonhuman primates can determine strangers' ranks by observing their interactions, and some evidence suggests that at least some cues to dominance, such as facial width-to-height ratio (fWHR), are also present in other primates. However, it is unknown whether they can determine strangers' rank simply by looking at their faces, rather than observing their interactions. If so, this would suggest selective pressure across the primates on both cues to dominance and the ability to detect those cues accurately. To address this, we examined the ability of male and female tufted capuchin monkeys (Sapajus [Cebus] apella) to categorize images of the faces of unknown conspecifics (Sapajus from different colonies) and humans (computer-generated and real) as dominant or nondominant based only on still images. Capuchins' categorization of unknown conspecific faces was consistent with fWHR, a cue to dominance, although there was a strong tendency to categorize strangers as dominant, particularly for males. This was true despite the continued correct categorization of known individuals. In addition, capuchins did not categorize human strangers in accordance with external pre-ratings of dominance by independent human raters, despite the availability of the same cue, fWHR. We consider these results in the context of capuchin socio-ecology and what they mean for the evolution of rapid decision-making in social contexts., (© 2024 Wiley Periodicals LLC.)

Social norms - rules governing which behaviours are deemed appropriate or inappropriate within a given community - are typically taken to be uniquely human. Recently, this position has been challenged by a number of philosophers, cognitive scientists, and ethologists, who have suggested that social norms may also be found in certain non-human animal communities. Such claims have elicited considerable scepticism from norm cognition researchers, who doubt that any non-human animals possess the psychological capacities necessary for normative cognition. However, there is little agreement among these researchers about what these psychological prerequisites are. This makes empirical study of animal social norms difficult, since it is not clear what we are looking for and thus what should count as behavioural evidence for the presence (or absence) of social norms in animals. To break this impasse, we offer an approach that moves beyond contested psychological criteria for social norms. This approach is inspired by the animal culture research program, which has made a similar shift away from heavily psychological definitions of 'culture' to become organised around a cluster of more empirically tractable concepts of culture. Here, we propose an analogous set of constructs built around the core notion of a normative regularity, which we define as a socially maintained pattern of behavioural conformity within a community. We suggest methods for studying potential normative regularities in wild and captive primates. We also discuss the broader scientific and philosophical implications of this research program with respect to questions of human uniqueness, animal welfare and conservation., (© 2024 The Authors. Biological Reviews published by John Wiley & Sons Ltd on behalf of Cambridge Philosophical Society.)

Many animals, including humans, must make decisions when outcomes involve risk and/or ambiguity. To explore the evolutionary roots of decision making when outcomes are unknown, we modified the Balloon Analogue Risk Task (BART) for use with tufted capuchin monkeys ( Sapajus [Cebus] apella ), creating the Primate Analogue Risk Task (PART). Using both the BART and the PART, we first compared human performance across the two tasks using analogous parameters. Humans' performance on the two tasks was positively correlated. Next, we tested capuchin monkeys' performance on the PART to assess their decision-making strategies in the context of ambiguity. Secondarily, although it was not the main goal of the study, this allowed us to look at species differences between capuchins' and humans' performance. Finally, we investigated the influence of prior experience on human and capuchin decision-making behavior. Neither capuchins nor humans behaved differently following an unsuccessful trial compared to a successful trial. We found individual differences in capuchin monkeys' choice behavior, though as a whole they demonstrated a pattern of reward maximization over time. Finally, as a species, capuchins had lower PART risk scores than humans. This paradigm presents a useful way to assess behavior in a context with uncertain outcomes using a comparative approach. (PsycInfo Database Record (c) 2024 APA, all rights reserved).

When making decisions, humans often strive to uphold objective, absolute standards, such as about what is small versus large, blue versus purple, or unfair versus fair, suggesting that our judgments should not be swayed by extraneous factors such as the sequence or frequency of events to be judged. Yet in previous research, when some items (e.g., threatening faces) became less frequent, humans responded by expanding their concept (of "threatening") to include more ambiguous stimuli. We assessed the origins of this perceptual frequency bias by testing 25 capuchins, seven rhesus monkeys, and 102 humans on a computer task in which they had to classify one circle at a time (pulled from a continuum of 50 circle sizes) as either small or large. Small and large circles initially appeared with equal probability but over time small circles either became less frequent, more frequent, or did not change in frequency. All three species showed changes in judgment, but contrary to predictions, they contracted, rather than expanded, their size judgments of the less frequent category. In other words, when small circles became rare, participants were more likely to judge ambiguous circles sizes as large (and vice versa). This may have been due to the immediate explicit feedback, as has recently been found in humans, and we consider possible mechanisms driving our participants' responses. These results suggest that humans' difficulties in maintaining absolute standards are shared with other animals. (PsycInfo Database Record (c) 2024 APA, all rights reserved).

Primatologist who brought animals and humans "a little closer".

The ability to quickly perceive and interpret threatening facial expressions from others is critical for successfully maintaining group cohesion in social nonhuman primate species. Rapid detection of threatening or negative stimuli in the environment compared to neutral stimuli, referred to as an attentional bias toward threat, is adaptive in that faster threat detection can lead to greater survival outcomes. However, the evolutionary roots of attentional bias formation toward social threat are not well understood. The present study investigated attentional biases toward social threat and the factors associated with them, including underlying hormonal mechanisms, in socially housed capuchin monkeys. Attentional biases were assessed using a dot-probe task that measured capuchins' latency to respond to a target using a joystick after viewing threatening or neutral conspecific or allospecific faces or nonface stimuli. In our first study, we examined how age, dominance status, sex, and cortisol level related to attentional biases. In our second study, we examined how manipulated oxytocin (OT) influenced attentional biases. Capuchin monkeys did not show attentional biases toward threatening faces or objects, but they showed attentional avoidance of scrambled familiar conspecific face stimuli. Cortisol and social rank were associated with attentional bias toward threat in the capuchin monkeys that participated in this study, which suggests that stress and dominance relate to attentional bias toward social threat. Manipulated OT increased attentional avoidance of scrambled familiar and unfamiliar face images, but not unscrambled faces or objects. Overall, we did not find compelling evidence of attentional biases toward social threat in capuchin monkeys. (PsycInfo Database Record (c) 2024 APA, all rights reserved).

Human cooperation can be facilitated by the ability to create a mental representation of one's own actions, as well as the actions of a partner, known as action co-representation. Even though other species also cooperate extensively, it is still unclear whether they have similar capacities. The Joint Simon task is a two-player task developed to investigate this action co-representation. We tested brown capuchin monkeys (Sapajus [Cebus] apella), a highly cooperative species, on a computerized Joint Simon task and found that, in line with previous research, the capuchins' performance was compatible with co-representation. However, a deeper exploration of the monkeys' responses showed that they, and potentially monkeys in previous studies, did not understand the control conditions, which precludes the interpretation of the results as a social phenomenon. Indeed, further testing to investigate alternative explanations demonstrated that our results were due to low-level cues, rather than action co-representation. This suggests that the Joint Simon task, at least in its current form, cannot determine whether non-human species co-represent their partner's role in joint tasks., (© 2024. The Author(s).)

Studies on coordination often present animals with the choice of either cooperating or remaining inactive; however, in nature, animals may also choose to act alone. This can be modeled with the Assurance game, an economic game that has recently been used to explore decision-making in primates. We investigated whether dyads of pet dogs coordinate in the Assurance game. Pairs were presented with two alternatives: they could individually solve an apparatus baited with a low-value reward ( Hare ) or they could coordinate to solve a cooperative apparatus baited with a high-value reward for each dog ( Stag ). All individuals matched their partner's choices, but after controlling for side bias, only four out of eleven dyads consistently coordinated on the payoff-dominant strategy ( Stag-Stag ). Thus, some dogs are capable of finding coordinated outcomes, as do primates, at least when their partner's actions are visible and coordination results in the biggest payoff for both individuals., Competing Interests: The authors declare no competing interests., (© 2023 The Authors.)

Economic models predict that rational decision makers' choices between a constant, "safe" option and a variable, "risky" option leading, on average, to the same payoff, should be random. However, a wealth of research has revealed that, when faced with risky decisions, both human and nonhuman animals deviate from economic rationality. According to the risk-sensitivity theory, individuals should prefer a safe option when they are in a positive energy state and a risky option when they are in a negative energy state. The abundance/risk hypothesis proposes that individuals should prefer risky options when diet quality exceeds their nutritional requirements. We tested how energy budget affects decision making under risk by presenting 22 capuchins belonging to two colonies (IT: N = 12, US: N = 10) with a risky choice task. Capuchins had to choose between a constant option (always four food items) and a variable option (one or seven food items with a 50% probability) in two conditions. In the Low-energy condition capuchins were tested before their main meal, whereas in the High-energy condition they were tested following a high-caloric meal. In neither colony did we find a significant difference between conditions, suggesting that the energy budget did not affect risk preferences. However, we found differences between colonies in their general response to risky choices: US capuchins were more risk-prone after selecting a safe option than a risky option and after selecting a bad (one food item) than a good (seven food items) risky option, whereas this did not hold true in IT capuchins. Furthermore, in the IT colony, males were more risk-prone under the High-energy condition compared to the Low-energy condition. Subtle differences in individual characteristics, management conditions, or stochastic founder effects may be implied, with relevant consequences for the outcomes of research on risky decision-making across laboratories., (© 2023 The Authors. American Journal of Primatology published by Wiley Periodicals LLC.)

Nonhuman primates exhibit sexual dimorphism in behavior, suggesting that there could be underlying differences in brain organization and function. Understanding this neuroanatomical variation is critical for enhancing our understanding of the evolution of sex differences in the human brain. Tufted capuchin monkeys (Sapajus [Cebus] apella) represent a phylogenetically diverse taxa of neotropical primates that converge on several behavioral characteristics with humans relevant to social organization, making them an important point of comparison for studying the evolution of sex differences in primates. While anatomical sex differences in gray matter have previously been found in capuchin monkeys, the current study investigates sex differences in white matter tracts. We carried out tract-based spatial statistical analysis on fractional anisotropy images of tufted capuchin monkeys (15 female, 5 male). We found that females showed significantly higher fractional anisotropy than males in regions of frontal-parietal white matter in the right cerebral hemisphere. Paralleling earlier findings in gray matter, male and female fractional anisotropy values in these regions were nonoverlapping. This complements prior work pointing toward capuchin sex differences in limbic circuitry and higher-order visual regions. We propose that these sex differences are related to the distinct socioecological niches occupied by male and female capuchins. Capuchin neuroanatomical sex differences appear to be more pronounced than in humans, which we suggest may relate to human adaptations for prolonged neurodevelopmental trajectories and increased plasticity., (© 2023 Wiley Periodicals LLC.)

Observed behavior can be the result of complex cognitive processes that are influenced by environmental factors, physiological process, and situational features. Pressure, a feature of a situation in which an individual's outcome is impacted by his or her own ability to perform, has been traditionally treated as a human-specific phenomenon and only recently have pressure-related deficits been considered in relation to other species. However, there are strong similarities in biological and cognitive systems among mammals (and beyond), and high-pressure situations are at least theoretically common in the wild. We hypothesize that other species are sensitive to pressure and that we can learn about the evolutionary trajectory of pressure responses by manipulating pressure experimentally in these other species. Recent literature indicates that, as in humans, pressure influences responses in non-human primates, with either deficits in ability to perform ("choking") or an ability to thrive when the stakes are high. Here, we synthesize the work to date on performance under pressure in humans and how hormones might be related to individual differences in responses. Then, we discuss why we would expect to see similar effects of pressure in non-humans and highlight the existing evidence for how other species respond. We argue that evidence suggests that other species respond to high-pressure contexts in similar ways as humans, and that responses to pressure are a critical missing piece of our understanding of cognition in human and non-human animals. Understanding pressure's effects could provide insight into individual variation in decision-making in comparative cognition and the evolution of human decision-making., (© 2023. The Author(s).)

Social interactions induce oxytocin release in many social species, suggesting that oxytocin is a critical part of social bonding among individuals. However, oxytocin also increases as a result of physical contact and stimulation, making it unclear which features of affiliative behaviors (for instance, social interaction or physical contact with a conspecific) drive the oxytocin increase observed after engaging in these behaviors. We attempted to tease this apart by studying the differential effect of social interaction, visual coordination with a conspecific, and physical stimulation during the fur-rubbing behavior of tufted capuchin monkeys (Sapajus [Cebus] apella), which often involves social contact with groupmates but is in some cases performed alone. We induced fur-rubbing by providing onions under three conditions: when capuchins had physical access to their social group and fur-rubbed in contact with groupmates (social condition), when capuchins were separated from their social group but could still see them fur-rub (visual coordination), and when capuchins were physically and visually separated from their groupmates (physical stimulation only). We assessed urinary oxytocin in these three conditions and compared them to a control condition in which apples were provided and no fur-rubbing was observed. Capuchins fur-rubbed for less time when they could not see their groupmates, but fur-rubbing increased urinary oxytocin above the control condition in all three fur-rubbing conditions equally, suggesting that the physical stimulation derived from fur-rubbing was the most important driver of oxytocin increase. These results support a model in which physical stimulation is an important factor in the relationship between oxytocin and at least some behaviors, suggesting that oxytocin increase alone is not necessarily indicative of a social influence on behavior. Future work is needed to determine the contexts in which social factors do impact oxytocin, and whether the downstream behaviors are the same for socially and nonsocially induced oxytocin release., (© 2023 Wiley Periodicals LLC.)

The origins of evolutionary games are rooted in both economics and animal behaviour, but economics has, until recently, focused primarily on humans. Although historically, specific games were used in targeted circumstances with non-human species (i.e. the Prisoner's Dilemma), experimental economics has been increasingly recognized as a valuable method for directly comparing both the outcomes of economic decisions and their underlying mechanisms across species, particularly in comparison with humans, thanks to the structured procedures that allow for them to be instantiated across a variety of animals. So far, results in non-human primates suggest that even when outcomes are shared, underlying proximate mechanisms can vary substantially. Intriguingly, in some contexts non-human primates more easily find a Nash equilibrium than do humans, possibly owing to their greater willingness to explore the parameter space, but humans excel at more complex outcomes, such as alternating between two Nash equilibria, even when deprived of language or instruction, suggesting potential mechanisms that humans have evolved to allow us to solve complex social problems. We consider what these results suggest about the evolution of economic decision-making and suggest future directions, in particular the need to expand taxonomic diversity, to expand this promising approach. This article is part of the theme issue 'Half a century of evolutionary games: a synthesis of theory, application and future directions'.

Humans stand out for their capacity to flexibly cooperate, possibly because they understand their partners' role. Researchers have explored if such understanding is unique to humans by assessing whether non-human species wait to manipulate a cooperative apparatus until a delayed partner arrives. If animals do wait, then it is assumed that they recognize the need for a partner. However, success in these tasks may be the result of social facilitation, while failure may be due to poor inhibitory control. Moreover, this approach does not test if animals take their partners' actions into account. Here we trained dogs to press a button simultaneously with their human partner. Afterwards, we tested them in several conditions to disentangle which elements of their partner's behaviour they take into account. Dogs waited to press the button until the delayed partner arrived, the button was available to the partner and the partner acted (pressed the button). We found no relationship between inhibitory control and success. We conclude that dogs are not merely reacting to the presence of their human partners, but are also taking their actions into account when coordinating with them.

Thinking about possibilities plays a critical role in the choices humans make throughout their lives. Despite this, the influence of individuals' ability to consider what is possible on culture has been largely overlooked. We propose that the ability to reason about future possibilities or prospective cognition, has consequences for cultural change, possibly facilitating the process of cumulative cultural evolution. In particular, by considering potential future costs and benefits of specific behaviours, prospective cognition may lead to a more flexible use of cultural behaviours. In species with limited planning abilities, this may lead to the development of cultures that promote behaviours with future benefits, circumventing this limitation. Here, we examine these ideas from a comparative perspective, considering the relationship between human and nonhuman assessments of future possibilities and their cultural capacity to invent new solutions and improve them over time. Given the methodological difficulties of assessing prospective cognition across species, we focus on planning, for which we have the most data in other species. Elucidating the role of prospective cognition in culture will help us understand the variability in when and how we see culture expressed, informing ongoing debates, such as that surrounding which social learning mechanisms underlie culture. This article is part of the theme issue 'Thinking about possibilities: mechanisms, ontogeny, functions and phylogeny'.

Primates of several species respond negatively to receiving less preferred rewards than a partner for completing the same task (inequity responses), either rejecting rewards or refusing to participate in the task when disadvantaged. This has been linked to cooperation, with species that cooperate frequently refusing to participate in inequity tasks (the 'cooperation hypothesis'). However, inequity is a social response, and previous research has involved dyads, precluding studying the effects of additional social partners. While dyads allow for tighter control in experimental settings, dyadic interactions in nature do not take place in a social vacuum, so understanding the role of the social context is needed to verify that the pattern of results supports the cooperation hypothesis. Here we focus on Bolivian squirrel monkeys, Saimiri boliviensis , a highly social species that does not generally cooperate and has not responded to inequity in previous dyadic research, although they do respond to receiving a lower reward than they expected. In the current study, we provide a more nuanced test by studying female Bolivian squirrel monkeys, the demographic most likely to cooperate in both field and laboratory contexts, in a more socially relevant group setting. For some reward values, females responded in both the inequity condition, rejecting less preferred rewards when they were disadvantaged relative to their social group, and a contrast condition, wherein all animals received a lower reward than they expected, making it difficult to disentangle contrast from inequity. As in capuchin monkeys, refusals increased when monkeys were to receive low-value rewards compared to medium-value rewards. These results suggest that the relationship between cooperation and inequity responses may be more nuanced than previously suggested, with demographic, social context and reward value potentially influencing outcomes even within species.

Visual attention to facial features is an important way that group-living primate species gain knowledge about others. However, where this attention is focused on the face is influenced by contextual and social features, and emerging evidence in Pan species suggests that oxytocin, a hormone involved in forming and maintaining affiliative bonds among members of the same group, influences social attention as measured by eye gaze. Specifically, bonobos tend to focus on conspecifics' eyes when viewing two-dimensional images, whereas chimpanzees focus more on the edges of the face. Moreover, exogenous oxytocin, which was hypothesized to increase eye contact in both species, instead enhanced this existing difference. We follow up on this to (1) determine the degree to which this Pan pattern generalizes across highly social, cooperative non-ape primates and (2) explore the impact of exogenously administered vs. endogenously released oxytocin in impacting this behavior. To do so, we tracked gaze direction on a computerized social categorization task using conspecific faces in tufted capuchin monkeys ( Sapajus [Cebus] apella ) after (1) exogenously administering intranasal oxytocin using a nebulizer or (2) inducing an endogenous increase in oxytocin using fur-rubbing, previously validated to increase oxytocin in capuchins. Overall, we did not find a general tendency in the capuchins to look toward the eyes or mouth, but we found that oxytocin was related to looking behavior toward these regions, albeit not in a straightforward way. Considering frequency of looking per trial, monkeys were more likely to look at the eye region in the fur-rubbing condition as compared to either the saline or exogenous oxytocin conditions. However, in terms of duration of looking during trials in which they did look at the eye region, monkeys spent significantly less time looking at the eyes in both oxytocin conditions as compared to the saline condition. These results suggest that oxytocin did not necessarily enhance eye looking in capuchins, which is consistent with the results from Pan species, and that endogenous and exogenous oxytocin may behave differently in their effect on how social attention is allocated., Competing Interests: The authors declare that the research was conducted in the absence of any commercial or financial relationships that could be construed as a potential conflict of interest., (Copyright © 2022 Sosnowski, Kano and Brosnan.)

The platyrrhine family Cebidae (capuchin and squirrel monkeys) exhibit among the largest primate encephalization quotients. Each cebid lineage is also characterized by notable lineage-specific traits, with capuchins showing striking similarities to Hominidae such as high sensorimotor intelligence with tool use, advanced cognitive abilities, and behavioral flexibility. Here, we take a comparative genomics approach, performing genome-wide tests for positive selection across five cebid branches, to gain insight into major periods of cebid adaptive evolution. We uncover candidate targets of selection across cebid evolutionary history that may underlie the emergence of lineage-specific traits. Our analyses highlight shifting and sustained selective pressures on genes related to brain development, longevity, reproduction, and morphology, including evidence for cumulative and diversifying neurobiological adaptations across cebid evolution. In addition to generating a high-quality reference genome assembly for robust capuchins, our results lend to a better understanding of the adaptive diversification of this distinctive primate clade.

Non-invasive health monitoring is advantageous for wild and captive primate populations because it reduces the need for traditional invasive techniques (i.e., anesthetization) that can be stressful and potentially harmful for individuals. The biomarker neopterin is an emerging tool in primatology to measure immune activation and immunosenescence, however, most neopterin studies have focused on catarrhine species with little comparative work examining neopterin and health in platyrrhines. To address this gap, we validated a commercially available enzyme-linked immunosorbent assay (ELISA) to measure urinary neopterin in two types of capuchin monkeys, a wild population of white-faced capuchins ( Cebus imitator ) and a socially housed captive colony of tufted capuchins ( Sapajus apella ). We analytically validated methods for measuring urinary neopterin in two capuchin populations and demonstrated that two commonly-used methods to control for urine concentration-creatinine and specific gravity (SG)-produced highly concordant results. We also biologically validated these methods by examining variation in neopterin levels based on environment (captive and wild) and age, and changes in levels associated with immune-response. We found that neopterin increased after immune perturbation (rabies vaccine booster), varied by environmental condition, and mirrored expected trends in immune system ontogeny. Our results improve understanding of the innate immune system in platyrrhine species and suggest neopterin may be useful for non-invasive health monitoring in both captive and wild primates., Competing Interests: The authors declare that the research was conducted in the absence of any commercial or financial relationships that could be construed as a potential conflict of interest., (Copyright © 2022 Lucore, Marshall, Brosnan and Benítez.)

Judges are typically tasked to consider sentencing benefits but not costs. Previous research finds that both laypeople and prosecutors discount the costs of incarceration when forming sentencing attitudes, raising important questions about whether professional judges show the same bias during sentencing. To test this, we used a vignette-based experiment in which Minnesota state judges ( N  = 87) reviewed a case summary about an aggravated robbery and imposed a hypothetical sentence. Using random assignment, half the participants received additional information about plausible negative consequences of incarceration. As predicted, our results revealed a mitigating effect of cost exposure on prison sentence term lengths. Critically, these findings support the conclusion that policies that increase transparency in sentencing costs could reduce sentence lengths, which has important economic and social ramifications., Competing Interests: The authors declare that the research was conducted in the absence of any commercial or financial relationships that could be construed as a potential conflict of interest., (Copyright © 2022 Aharoni, Kleider-Offutt, Brosnan and Hoffman.)

Humans often experience striking performance deficits when their outcomes are determined by their own performance, colloquially referred to as "choking under pressure." Physiological stress responses that have been linked to both choking and thriving are well-conserved in primates, but it is unknown whether other primates experience similar effects of pressure. Understanding whether this occurs and, if so, its physiological correlates, will help clarify the evolution and proximate causes of choking in humans. To address this, we trained capuchin monkeys on a computer game that had clearly denoted high- and low-pressure trials, then tested them on trials with the same signals of high pressure, but no difference in task difficulty. Monkeys significantly varied in whether they performed worse or better on high-pressure testing trials and performance improved as monkeys gained experience with performing under pressure. Baseline levels of cortisol were significantly negatively related to performance on high-pressure trials as compared to low-pressure trials. Taken together, this indicates that less experience with pressure may interact with long-term stress to produce choking behavior in early sessions of a task. Our results suggest that performance deficits (or improvements) under pressure are not solely due to human specific factors but are rooted in evolutionarily conserved biological factors., (© 2022. The Author(s).)

Collective decision-making is a widespread phenomenon across organisms. Studying how animal societies make group decisions to the mutual benefit of group members, while avoiding exploitation by cheaters, can provide unique insights into the underlying cognitive mechanisms. As a step toward dissecting the proximate mechanisms that underpin collective decision-making across animals, we developed an agent-based model of antipredatory alarm signaling and mobbing during predator-prey encounters. Such collective behaviors occur in response to physical threats in many distantly related species with vastly different cognitive abilities, making it a broadly important model behavior. We systematically assessed under which quantitative contexts potential prey benefit from three basic strategies: predator detection, signaling about the predator (e.g., alarm calling), and retreating from vs. approaching the predator. Collective signaling increased survival rates over individual predator detection in several scenarios. Signaling sometimes led to fewer prey detecting the predator but this effect disappeared when prey animals that had seen the predator both signaled and approached it, as in mobbing. Critically, our results highlight that collective decision-making in response to a threat can emerge from simple rules without needing a central leader or needing to be under conscious control., (Copyright © 2021 Elsevier B.V. All rights reserved.)

Prosecutors can influence judges' sentencing decisions by the sentencing recommendations they make-but prosecutors are insulated from the costs of those sentences, which critics have described as a correctional "free lunch." In a nationally distributed survey experiment, we show that when a sample of ( n =178) professional prosecutors were insulated from sentencing cost information, their prison sentence recommendations were nearly one-third lengthier than sentences rendered following exposure to direct cost information. Exposure to a fiscally equivalent benefit of incarceration did not impact sentencing recommendations, as predicted. This pattern suggests that prosecutors implicitly value incorporating sentencing costs but selectively neglect them unless they are made explicit. These findings highlight a likely but previously unrecognized contributor to mass incarceration and identify a potential way to remediate it., Competing Interests: The authors declare that the research was conducted in the absence of any commercial or financial relationships that could be construed as a potential conflict of interest., (Copyright © 2021 Aharoni, Kleider-Offutt and Brosnan.)

Our understanding of animals' affective processing is notably limited compared to the wealth of research on humans, largely due to difficulties in measurement. Moreover, despite a recent increase in the understanding of the interaction between affect and cognition in animals, most research has focused on negative affect, with the result that we continue to know little about the effects of positive affect. In this study, we tested 15 adult capuchin monkeys ( Sapajus [ Cebus ] apella ) using a novel methodology that took advantage of capuchins' species-typical behavior to engineer both a positive and negative experience, using the same apparatus to minimize extraneous impacts. Following a positive or negative experience (that presumably induced positive and negative affect, respectively), or a control with no manipulation, we assessed subjects' performance on a cognitive task, a computerized delayed match-to-sample. As predicted, behavior following the negative condition suggested a negative affective state, with increased rates of scratching (commonly used as an indicator of stress in nonhuman primates) compared to both the positive and control conditions. Cognitive performance was also impaired in the negative condition compared to the other two. Contrary to predictions, however, the positive condition did not have a facilitative effect on cognitive performance, but behavioral results indicate that we may not have induced a truly positive affective state. Although we add to evidence that a negative experience can influence subsequent behavior and cognitive performance in nonhuman primates, our work highlights our lack of knowledge about the impact of positive affect, if any, on behavior and cognition. (PsycInfo Database Record (c) 2021 APA, all rights reserved).

Unequal outcomes disrupt cooperation in some situations, but this has not been tested in the context of coordination in economic games. To explore this, we tested brown capuchins (Sapajus [Cebus] apella) on a manual version of the Stag Hunt (or Assurance) Game, in which individuals sequentially chose between two options, Stag or Hare, and were rewarded according to their choices and that of their partner. Typically, coordination on Stag results in an equal highest payout, whereas coordinating on Hare results in a guaranteed equal but lower payoff and uncoordinated play results in the lowest payoff when playing Stag. We varied this structure such that one capuchin received double the rewards for the coordinated Stag outcome; thus, it was still both animals' best option, but no longer equally rewarding. Despite the inequality, capuchins coordinated on Stag in 78% of trials, and neither payoff structure nor their partner's choice impacted their decision. Additionally, there was no relationship between self-scratching, a measure of stress in capuchins, and choices. After completing the study, we discovered our reward, cheerios, was sufficiently valuable that in another study, capuchins never refused it, so post hoc we repeated the study using a lower value reward, banana flavored pellets. Capuchins completed only 26% of the pellet trials (compared to 98% with cheerios), constraining our ability to interpret the results, but nonetheless the monkeys showed a decrease in preference for Stag, particularly when they received fewer rewards for the coordinated Stag outcome. These results reinforce capuchins' ability to find coordinated outcomes in the Stag Hunt game, but more work is needed to determine whether the monkeys did not mind the inequality or were unwilling to sacrifice a highly preferred food to rectify it. In either case, researchers should carefully consider the impact of their chosen rewards on subjects' choices., (© 2021 The Authors. American Journal of Primatology published by Wiley Periodicals LLC.)

Although individuals in some species refuse foods they normally accept if their partner receives a more preferred one, this is not true across all species. The cooperation hypothesis proposes that this species-level variability evolved because inequity aversion is a mechanism to identify situations in which cooperation is not paying off, and that species regularly observed cooperating should be more likely to be averse to inequity. To rule out other potential explanations of inequity aversion, we need to test the converse as well: species rarely observed cooperating, especially those phylogenetically close to more cooperative species, should be less likely to be inequity averse. To this end, we tested eight zoo-housed Western lowland gorillas (Gorilla gorilla gorilla) on a token exchange task in which subjects received either the same food reward or a less-preferred reward for the same or more effort than their partner, recording both refusals to participate in the exchange and refusals to accept the reward. Supporting the cooperation hypothesis, even with procedural differences across sessions, gorillas were significantly more likely to refuse in all conditions in which they received a low-value food reward after completing an exchange, regardless of what their partner received, suggesting that gorillas were not inequity averse, but instead would not work for a low-value reward. Additionally, gorillas were more likely to refuse later in the session; while the pattern of refusals remained unchanged after accounting for this, this suggests that species should be tested on as many trials as is practical., (© 2021 Wiley Periodicals LLC.)

Cumulative cultural evolution (CCE), the improvement of cultural traits over generations via social transmission, is widely believed to be unique to humans. The capacity to build upon others' knowledge, technologies, and skills has produced the most diverse and sophisticated technological repertoire in the animal kingdom. Yet, inconsistency in both the definitions and criteria used to determine CCE and the methodology used to examine it across studies may be hindering our ability to determine which aspects are unique to humans. Issues regarding how improvement is defined and measured and whether some criteria are empirically testable are of increasing concern to the field. In this article, we critically assess the progress made in the field and current points of debate from conceptual and methodological perspectives. We discuss how inconsistency in definitions is detrimental to our ability to document potential evidence of CCE to nonhuman animals. We build on Mesoudi and Thornton's (2018) recently described core and extended CCE criteria to make specific recommendations about, from a comparative lens, which criteria should be used as evidence of CCE. We evaluate existing data from both wild and captive studies of nonhuman animals using these suggestions. We finish by discussing issues currently faced by researchers studying CCE in nonhuman animals, particularly nonhuman primates, and provide suggestions that may overcome these concerns and move the field forward. (PsycInfo Database Record (c) 2021 APA, all rights reserved).

As a relatively young discipline, primatology is lucky to still have many of its pioneering researchers active in the field. At the 2019 American Society of Primatology meeting, I convened the inaugural Pioneers in Primatology symposium to bring together a selection of these researchers to reflect on their careers, with a focus on the stories that defined their work and their thoughts on the important future directions for the field. After the symposium, Karen Strier and I decided that these pioneers' contributions merited preservation, and this special issue is our effort to do so., (© 2021 Wiley Periodicals LLC.)

Large-scale studies of individual differences in innovative behavior among nonhuman animals are rare because of logistical difficulties associated with obtaining observational data on a large number of innovative individuals across multiple locations. Here, we take a different approach, using observer ratings to study individual differences in innovative behavior in 127 brown capuchin monkeys (Sapajus [Cebus] sp.) from 15 social groups and 7 facilities. Capuchins were reliably rated by 1 to 7 raters (mean 3.2 ± 1.6 raters/monkey) on a 7-point Likert scale for levels of innovative behavior, task motivation, sociality, and dominance. In a subsample, we demonstrate these ratings are valid: Rated innovation predicted performance on a learning task, rated motivation predicted participation in the task, rated dominance predicted social rank based on win/loss aggressive outcomes, and rated sociality predicted the time that monkeys spent in proximity to others. Across all 127 capuchins, individuals that were rated as being more innovative were significantly younger, more social, and more motivated to engage in tasks. Age, sociality, and task motivation all had independent effects on innovativeness, whereas sex, dominance, and group size were nonsignificant. Our findings are consistent with long-term behavioral observations of innovation in wild white-faced capuchins. Observer ratings may, therefore, be a valid tool for studies of animal innovation. (PsycInfo Database Record (c) 2021 APA, all rights reserved).

In the past decade, there has been a surge of interest in using games derived from experimental economics to test decision-making behaviour across species. In most cases, researchers are using the games as a tool, for instance, to understand what factors influence decision-making, how decision-making differs across species or contexts, or to ask broader questions about species' propensities to cooperate or compete. These games have been quite successful in this regard. To what degree, however, do these games tap into species' economic decision-making? For the purpose of understanding the evolution of economic systems in humans, this is the key question. To study this, we can break economic decision-making down into smaller components, each of which is a potential step in the evolution of human economic behaviour. We can then use data from economic games, which are simplified, highly structured models of decision-making and therefore ideal for the comparative approach, to directly compare these components across species and contexts, as well as in relation to more naturalistic behaviours, to better understand the evolution of economic behaviour and the social and ecological contexts that influenced it. The comparative approach has successfully informed us about the evolution of other complex traits, such as language and morality, and should help us more deeply understand why and how human economic systems evolved. This article is part of the theme issue 'Existence and prevalence of economic behaviours among non-human primates'.

This study reports an analysis of 20 T1-weighted magnetic resonance imaging scans from tufted capuchin monkeys (5 male, 15 female). We carried out a data-driven, whole-brain volumetric analysis on regional gray matter anatomy using voxel-based morphometry. This revealed that males showed statistically significant expansion of a region of the hypothalamus, while females showed significant expansion in a distributed set of regions, including the cerebellum, early visual cortex, and higher-order visual regions spanning occipital and temporal cortex. In order to elucidate the network connectivity of these regions, we employed probabilistic tractography on diffusion tensor imaging data. This showed that the female-enlarged regions connect with distributed association networks across the brain. Notably, this contrasts with rodent studies, where sex differences are focused in deep, ancestral limbic regions involved in the control of reproductive behavior. Additionally, in our data set, for several regions, male and female volumetric measures were completely nonoverlapping. This contrasts with human studies, where sex differences in cortical regions have been reported but are characterized by overlapping rather than divergent male and female values. We suggest that these results can be understood in the context of the different lifetime experiences of males and females, which may produce increased experience-dependent cortical plasticity in capuchins compared to rodents, and in humans compared to capuchins., (© 2020 Wiley Periodicals LLC.)

A goal of the comparative approach is to test a variety of species on the same task. Here, we examined whether the factors that helped capuchin monkeys improve their performance in a dichotomous choice task would generalize to three other primate species: orangutans, gorillas, and drill monkeys. In this task, subjects have access to two options, each resulting in an identical food, but one (the ephemeral option) is only available if it is chosen first, whereas the other one (the permanent option) is always available. Therefore, the food-maximizing solution is to choose the ephemeral option first, followed by the permanent option for an additional reward. On the original version (plate task), the options were discriminated by the color and pattern of the plates holding the food, while on two subsequent versions we used altered cues that we predicted would improve performance: (1) the color of the foods themselves (color task), which we hypothesized was relevant to primates, who choose foods rather than substrates on which foods are found when foraging, and (2) patterned cups covering the foods (cup task), which we hypothesized would help primates avoid the prepotent response associated with visible food. Like capuchins, all three species initially failed to solve the plate task. However, while orangutans improved their performance from the plate to the color task, they did not for the cup task, and only a few gorillas and no drills succeeded in either task. Unfortunately, our ability to interpret these data was obscured by differences in the subjects' level of experience with cognitive testing and practical constraints that precluded the use of completely identical procedures across species. Nonetheless, we consider what these results can tell us, and discuss the value of conducting studies across multiple sites despite unavoidable differences., (© 2020 Wiley Periodicals LLC.)

Understanding the distribution of and mechanisms underlying prosocial behavior in other species has been hampered by methodological differences among studies, questions about animals' interpretations of the designs, and low phylogenetic diversity. Brucks and von Bayern's recent report that African gray parrots are proactively prosocial reiterates the importance of social tolerance and suggests a role for perspective taking, but additional work using identical procedures across taxa is needed to verify the cognitive mechanisms underpinning prosocial behavior.

Human decision-making is often swayed by irrecoverable investments even though it should only be based on future-and not past-costs and benefits. Although this sunk cost effect is widely documented and can lead to devastating losses, the underlying psychological mechanisms are unclear. To tease apart possible explanations through a comparative approach, we assessed capuchin and rhesus monkeys' susceptibility to sunk costs in a psychomotor task. Monkeys needed to track a moving target with a joystick-controlled cursor for variable durations. They could stop at any time, ending the trial without reward. To minimize the work required for a reward, monkeys should have always persisted for at least 1 s, but should have abandoned the trial if that did not yield a reward. Capuchin monkeys and especially rhesus macaques persisted to trial completion even when it was suboptimal, and were more likely to complete the trial the longer they had already tracked the target. These effects were less pronounced, although still present, when the change in expected tracking duration was signalled visually. These results show that sunk cost effects can arise in the absence of human-unique factors and may emerge, in part, because persisting can resolve uncertainty.

Practical and ethical constraints limit our ability to experimentally test socioecological theory in wild primates. We took an alternate approach to model this, allowing groups of humans to interact in a virtual world in which they had to find food and interact with both ingroup and outgroup avatars to earn rewards. We altered ratios and distributions of high- and low-value foods to test the hypothesis that hominoids vary with regards to social cohesion and intergroup tolerance due to their feeding ecology. We found larger nesting clusters and decreased attacks on outgroup competitors in the Bonobo condition versus the Chimpanzee condition, suggesting a significant effect of feeding competition alone on social structure. We also demonstrate that virtual worlds are a robust mechanism for testing hypotheses that are impossible to study in the wild.

Anthropomorphism is the attribution of human-like capacities and traits to non-human entities. Anthropomorphism is ubiquitous in everyday life and in scientific domains, operating both implicitly and explicitly as a function of the human lens through which we view the world. A rich history of work in psychology, animal behavior, cognitive science, and philosophy has highlighted the negative and, to a lesser degree, the positive implications of anthropomorphism. In this article, we aim to provide a nuanced perspective of how anthropomorphism impacts the work of comparative affective science. Specifically, we discuss three domains of empirical inquiry in which lessons can be drawn about the benefits and pitfalls of anthropomorphism: responses to death, inequity aversion, and prosocial behavior. On balance, we advocate a mindful approach to anthropomorphizing in comparative affective science, and comparative science more generally., Competing Interests: Declaration of Competing Interest None., (Copyright © 2020 Elsevier Ltd. All rights reserved.)

Do people punish more than they would if the decision costs were more transparent? In two Internet-based vignette experiments, we tested whether juvenile sentencing recommendations among U.S. adults are responsive to variation in the salience of the taxpayer costs and public safety benefits of incarceration. Using a 2 Cost (present vs. absent) x 2 Benefit (present vs. absent) factorial design, Experiment 1 (N = 234) found that exposure to information about the direct costs of incarcerating the juvenile offender reduced sentencing recommendations by about 28%, but exposure to the public safety benefits had no effect on sentences. Experiment 2 (N = 301) manipulated cost-benefit salience by asking participants to generate their own list of costs of incarceration, benefits of incarceration, or an affectively neutral, unrelated word list. Results revealed a similar selective effect whereby sentencing recommendations were reduced in the cost condition relative to the benefits and control conditions, but sentences in the benefit condition did not differ from the control. This combined pattern suggests that laypeople selectively neglect to factor cost considerations into these judgments, thereby inflating their support for punishment, unless those costs are made salient. These findings contribute to the debate on transparency in sentencing., Competing Interests: The authors have declared that no competing interests exist.

We review and analyze evidence for an evolutionary rooting of human economic behaviors and organization in non-human primates. Rather than focusing on the direct application of economic models that a priori account for animal decision behavior, we adopt an inductive definition of economic behavior in terms of the contribution of individual cognitive capacities to the provision of resources within an exchange structure. We spell out to what extent non-human primates' individual and strategic decision behaviors are shared with humans. We focus on the ability to trade, through barter or token-mediated exchanges, as a landmark of an economic system among members of the same species. It is an open question why only humans have reached a high level of economic sophistication. While primates have many of the necessary cognitive abilities (symbolic and computational) in isolation, one plausible issue we identify is the limits in exerting cognitive control to combine several sources of information. The difference between human and non-human primates' economies might well then be in degree rather than kind., (Copyright © 2019 Elsevier Ltd. All rights reserved.)

A central premise of the science of comparative affect is that we can best learn about the causes and consequences of affect by comparing affective phenomena across a variety of species, including humans. We take as a given that affect is widely shared across animals, but a key challenge is to accurately represent each species' affective experience. A common approach in the comparative study of behavior and cognition is to develop standardized experimental paradigms that can be used across species, with the assumption that if the same task is being used, we can directly compare behavioral responses. This experimental approach rests on two underlying assumptions: first, that different species' perception of and affective response to these paradigms are the same; and second, that behavioral and physiological (including endocrine and neural) responses to these paradigms are homologous; if either of these assumptions is not true, then the comparison becomes much less straightforward. Our goal in the present paper is to summarize the dominant paradigms that have been used for such comparative research, with a particular focus on paradigms common in the cooperation literature, and to critically discuss dominant assumptions about what affective states these tasks can or should measure. We then consider the advantages and drawbacks of this experimental method, and consider alternatives that may improve our understanding. We hope that this will help scholars recognize and avoid pitfalls inherent in studying affect, and stimulate them to create novel, ecologically relevant paradigms for examining affect across the animal kingdom., (Copyright © 2019 The Authors. Published by Elsevier Ltd.. All rights reserved.)

Learned rules help us accurately solve many problems, but by blindly following a strategy, we sometimes fail to find more efficient alternatives. Previous research found that humans are more susceptible to this "cognitive set" bias than other primates in a nonverbal computer task. We modified the task to test one hypothesis for this difference, that working memory influences the advantage of taking a shortcut. During training, 60 humans, 7 rhesus macaques, and 22 capuchin monkeys learned to select three icons in sequence. They then completed 96 baseline trials, in which only this learned rule could be used, and 96 probe trials, in which they could also immediately select the final icon. Rhesus and capuchin monkeys took this shortcut significantly more often than humans. Humans used the shortcut more in this new, easier task than in previous work, but started using it significantly later than the monkeys. Some participants of each species also used an intermediate strategy; they began the learned rule but switched to the shortcut after selecting the first item in the sequence. We suggest that these species differences arise from differences in rule encoding and in the relative efficiency of exploiting a familiar strategy versus exploring alternatives.

Serotonin is a neurotransmitter that plays an important role in regulating behavior and personality in humans and other mammals. Polymorphisms in genes coding for the serotonin receptor subtype 1A (HTR1A), the serotonin transporter (SLC6A4), and the serotonin degrading enzyme monoamine oxidase A (MAOA) are associated with anxiety, impulsivity, and neurotic personality in humans. In primates, previous research has largely focused on SLC6A4 and MAOA, with few studies investigating the role of HTR1A polymorphic variation on behavior. Here, we examined variation in the coding region of HTR1A across apes, and genotyped polymorphic coding variation in a sample of 214 chimpanzees with matched measures of personality and behavior. We found evidence for positive selection at three amino acid substitution sites, one in chimpanzees-bonobos (Thr26Ser), one in humans (Phe33Val), and one in orangutans (Ala274Gly). Investigation of the HTR1A coding region in chimpanzees revealed a polymorphic site, where a C/A single nucleotide polymorphism changes a proline to a glutamine in the amino acid sequence (Pro248Gln). The substitution is located in the third intracellular loop of the receptor, a region important for serotonin signal transduction. The derived variant is the major allele in this population (frequency 0.67), and is associated with a reduction in anxiety, decreased rates of male agonistic behavior, and an increase in socio-positive behavior. These results are the first evidence that the HTR1A gene may be involved in regulating social behavior in chimpanzees and encourage further systematic investigation of polymorphic variation in other primate populations with corresponding data on behavior., (© The Author(s) 2019. Published by Oxford University Press on behalf of the Society for Molecular Biology and Evolution. All rights reserved. For permissions, please e-mail: journals.permissions@oup.com.)

Comparative approaches to experimental economics have shed light on the evolution of social decision-making across a range of primate species, including humans. Here we replicate our previous work looking at six pairs of capuchin monkeys' (Sapajus [Cebus] apella) responses to scenarios requiring both coordination (Assurance Game) and anti-coordination (Hawk-Dove Game). This then provides a foundation for assessing their responses to two additional games, one with a scenario of beneficial cooperation with a temptation to defect (Prisoner's Dilemma) and one with an environment requiring changing strategies within short temporal proximity (Alternating Economic Game). We additionally explored the effects of exogenous oxytocin on decision-making. Oxytocin did not affect decisions in any of our games. Results from the first two games largely replicated our previous findings. Responses to the Prisoner's Dilemma were more varied than was seen in previous games, with pairs respectively cooperating, defecting, and failing to establish stable strategies. Such variability indicates that this game may be a good assay for individual differences in social decision-making. Finally, capuchins were able to flexibly switch between their previously established strategies within each of the different games, even when the games were presented within the same session, requiring strategy adjustments within short temporal proximity. These results build on earlier findings showing that capuchins can alter decision-making strategies as the context demands, which is likely essential for decision-making in naturally occurring contexts., (© 2019 Wiley Periodicals, Inc.)

Games from experimental economics have provided insights into the evolutionary roots of social decision making in primates and other species. Multiple primate species' abilities to cooperate, coordinate and anti-coordinate have been tested utilizing variants of these simple games. Past research, however, has focused on species known to cooperate and coordinate in the wild. To begin to address the degree to which cooperation and coordination may be a general ability that manifests in specific contexts, the present study assessed the decisions of squirrel monkeys ( Saimiri boliviensis; N = 10), a species not known for their cooperative behavior in these games. Pairs of monkeys were presented with the Assurance Game (a coordination game), the Hawk-Dove Game (an anti-coordination game) and the Prisoner's Dilemma (a cooperation game with a temptation to defect). We then compared squirrel monkeys' performance to existing data on capuchin monkeys ( Sapajus [Cebus] apella ), a closely related species that routinely cooperates, to determine what, if any, differences in decision making emerged. Some pairs of both species found the payoff-dominant Nash Equilibrium (NE) in the coordination game, but failed to find the NE in subsequent games. Our results suggest that, like capuchins, squirrel monkeys coordinate their behavior with others, suggesting that such mutual outcomes occur in at least some contexts, even in species that do not routinely cooperate.

The capacity for planning in nonhuman species has long been an interest of many comparative and cognitive psychologists. There is now considerable evidence that at least great apes show both motor planning and planning for future needs in various contexts and modalities. Few studies, however, have investigated planning ability in a monkey species, and most of these exceptions have used computerized procedures. To gain a broader view, in the current study, we tested capuchin monkeys' capacity for motor planning using the "paddle-box" apparatus, a manual maze task originally designed for testing planning skills in apes (Tecwyn, Thorpe, & Chappell, 2013). This consisted of a box containing different levels of paddles that subjects rotated to drop food into a tube that delivered it to the subject. To evaluate subjects' degree of skill in the task, the initial locations of the food and the final goal (i.e., the "paths" the food could take) were chosen according to the probability of retrieving the reward, starting with those that gave the highest probability of success and moving sequentially to those with the lowest probability. Most subjects solved all levels of difficulty in the task, and capuchin monkeys succeeded as a group in a generalization test with novel paths. These findings demonstrate that some monkeys, like apes, show planning in different contexts and modalities. (PsycINFO Database Record (c) 2019 APA, all rights reserved).

Games derived from experimental economics can be used to directly compare decision-making behavior across primate species, including humans. For example, the use of coordination games, such as the Assurance game, has shown that a variety of primate species can coordinate; however, the mechanism by which they do so appears to differ across species. Recently, these games have been extended to explore anti-coordination and cooperation in monkeys, with evidence that they play the Nash equilibria in sequential games in these other contexts. In the current paper, we use the same methods to explore chimpanzees' behavior in the Assurance Game; an anti-coordination game, the Hawk Dove game; and a cooperation game with a temptation to defect, the Prisoner's Dilemma game. We predicted that they would consistently play the Nash equilibria, as do the monkeys, and that, as in previous work, the subjects' level of experience with cognitive experiments would impact performance. Surprisingly, few of our pairs consistently played the same outcome (i.e., no statistically significant preferences), although those who did showed evidence consistent with Nash equilibria play, the same pattern seen more consistently in the monkeys. We consider reasons for their inconsistent performance; for instance, perhaps it was due to lack of interest in a task that rewarded them almost every trial no matter what option they chose, although this does not explain why they were inconsistent when the monkeys were not. A second goal of our study was to ascertain the effects of exogenous oxytocin in their decision making in one population. In line with recent work showing complex effects of oxytocin on social behavior, we found no effect on subjects' outcomes. We consider possible explanations for this as well.