Testosterone and cortisol jointly regulate dominance: Evidence for a dual-hormone hypothesis

Traditional theories propose that testosterone should increase dominance and other status-seeking behaviors, but empirical support has been inconsistent. The present research tested the hypothesis that testosterone's effect on dominance depends on cortisol, a glucocorticoid hormone implicated in psychological stress and social avoidance. In the domains of leadership (Study 1, mixed-sex sample) and competition (Study 2, male-only sample), testosterone was positively related to dominance, but only in individuals with low cortisol. In individuals with high cortisol, the relation between testosterone and dominance was blocked (Study 1) or reversed (Study 2). Study 2 further showed that these hormonal effects on dominance were especially likely to occur after social threat (social defeat). The present studies provide the first empirical support for the claim that the neuroendocrine reproductive (HPG) and stress (HPA) axes interact to regulate dominance. Because dominance is related to gaining and maintaining high status positions in social hierarchies, the findings suggest that only when cortisol is low should higher testosterone encourage higher status. When cortisol is high, higher testosterone may actually decrease dominance and in turn motivate lower status.     

Subclinical Primary Psychopathy,but Not Physical Formidability or Attractiveness,Predicts Conversational Dominance in a Zero-Acquaintance Situation

The determinants of conversational dominance are not well understood. We used videotaped triadic interactions among unacquainted same-sex American college students to test predictions drawn from the theoretical distinction between dominance and prestige as modes of human status competition. Specifically, we investigated the effects of physical formidability, facial attractiveness, social status, and self-reported subclinical psychopathy on quantitative (proportion of words produced), participatory (interruptions produced and sustained), and sequential (topic control) dominance. No measure of physical formidability or attractiveness was associated with any form of conversational dominance, suggesting that the characteristics of our study population or experimental frame may have moderated their role in dominance dynamics. Primary psychopathy was positively associated with quantitative dominance and (marginally) overall triad talkativeness, and negatively associated (in men) with affect word use, whereas secondary psychopathy was unrelated to conversational dominance. The two psychopathy factors had significant opposing effects on quantitative dominance in a multivariate model. These latter findings suggest that glibness in primary psychopathy may function to elicit exploitable information from others in a relationally mobile society.     

Testosterone and cortisol jointly regulate dominance: Evidence for a dual-hormone hypothesis

Traditional theories propose that testosterone should increase dominance and other status-seeking behaviors, but empirical support has been inconsistent. The present research tested the hypothesis that testosterone's effect on dominance depends on cortisol, a glucocorticoid hormone implicated in psychological stress and social avoidance. In the domains of leadership (Study 1, mixed-sex sample) and competition (Study 2, male-only sample), testosterone was positively related to dominance, but only in individuals with low cortisol. In individuals with high cortisol, the relation between testosterone and dominance was blocked (Study 1) or reversed (Study 2). Study 2 further showed that these hormonal effects on dominance were especially likely to occur after social threat (social defeat). The present studies provide the first empirical support for the claim that the neuroendocrine reproductive (HPG) and stress (HPA) axes interact to regulate dominance. Because dominance is related to gaining and maintaining high status positions in social hierarchies, the findings suggest that only when cortisol is low should higher testosterone encourage higher status. When cortisol is high, higher testosterone may actually decrease dominance and in turn motivate lower status.     

Male social status and its predictors among Garisakang forager-horticulturalists of lowland Papua New Guinea

Social hierarchy exists in all human societies, yet the characteristics important for achieving high social status may differ cross-culturally. Studies among Western industrialized populations often use indices of socioeconomic status (SES) to determine individual social position. Conversely, studies among small-scale societies typically use locally relevant traits. Cross-cultural applicability and the relationships between these two strategies have rarely been investigated. Studies limited to industrialized societies demonstrate that low social status is associated with poor health and elevated levels of chronic stress. It remains unclear, however, if and to what extent this relationship is a recent phenomenon in industrialized Western society or a universal human characteristic that has likely been present throughout much of our recent evolutionary history. In this study, we investigate relationships between various SES and locally relevant measures of male social status, age, body-mass index (BMI), and levels of waking salivary cortisol among relatively egalitarian Garisakang forager-horticulturalists of lowland Papua New Guinea. We employ a photo-ranking method using 32 raters (16 women, 16 men) to evaluate 15 measures of social hierarchy in men (N = 32). These measures target locally relevant traits considered important for male social status (e.g., sociability, hunting skills, fighting ability, and community influence). Using principal component analysis (PCA), we extracted two components labeled as Dominance-Respect and Prosociality-Skills. Models investigating predictors of social status demonstrate that age was significantly and positively related to both Dominance-Respect and Prosociality-Skills while BMI was positively related to only Dominance-Respect. Neither Dominance-Respect nor Prosociality-Skills predicted salivary cortisol levels. However, the SES measure of Income was significantly and positively related to cortisol levels, suggesting that men with higher income experience higher levels of stress. This finding may be explained by increased psychological stress accompanying market integration among the Garisakang or, alternatively, by culturally specific expectations for resource sharing and fear of conflict. These findings emphasize the usefulness of diverse social status measures in anthropological research and stress the need for broadened perspectives regarding the causes and consequences of social status among small-scale populations.     

Dominance and prestige as differential predictors of aggression and testosterone levels in men

The relationship between self-esteem and aggression has yielded mixed results and generated much recent debate in the social psychology literature. Based on an evolutionary-psychological theory of self-esteem, Kirkpatrick et al. [Kirkpatrick, L. A., Waugh, C. E., Valencia, A., Webster, G., 2002. The functional domain-specificity of self-esteem and the differential prediction of aggression. Journal of Personality and Social Psychology 825, 756–767] showed that multiple, functionally distinct self-esteem mechanisms predict aggression differentially: e.g., “self-perceived superiority” is positively related, and “social inclusion” inversely related, to behavioral aggression. The present study extends this research by further differentiating two distinct forms of “superiority,” dominance and prestige [Henrich, J., Gil-White, F. J. 2001. The evolution of prestige: Freely conferred deference as a mechanism for enhancing the benefits of cultural transmission. Evolution and Human Behavior 22, 165–196], in the prediction of aggression in men and women and testosterone levels (measured in saliva samples) in men. Dominance was positively related, but prestige was either unrelated or inversely related, to self-report aggression measures. Dominance was unrelated but prestige inversely related to testosterone levels in men, perhaps suggesting a method of testosterone inhibition in individuals attaining prestige-based superiority. In addition to contributing to the growing literature on the aggression–self-esteem link, the results provide validation for the prestige–dominance distinction and support, but also suggest an important refinement to, a theory of self-esteem as a collection of functionally distinct adaptations.     

Evidence for a nonverbal expression of triumph

An expression signaling victory in agonistic encounters may help to establish status and dominance in a hierarchy. We investigated the possibility that an expression of triumph is such a signal. In Study 1, spontaneous expressions of athletes who had just won a medal at the 2004 Olympic Games were shown to three groups of observers who judged the emotion portrayed. All were given a set of common response alternatives that included Anger, Contempt, Disgust, Fear, Joy, Sadness, Surprise, Neutral, and Other. Group 1 was also given Pride, Group 2 Triumph, and Group 3 both Pride and Triumph. Group 1 labeled some expressions as pride, but Group 2 labeled all the expressions as triumph. For Group 3, some expressions were consistently labeled as triumph, and others as pride. Behavioral coding indicated that the expressions labeled triumph were distinct from other expressions. In Study 2, we replicated this finding with a judgment task involving more positive emotion choices to eliminate the possibility that the previous findings occurred because of a limited number of positive emotion labels. In Study 3, we replicated the findings again in two groups of observers from South Korea using two different judgment tasks. These findings suggest that triumph may have a unique nonverbal expression.     

Direct and indirect selection on flowering time,water‐use efficiency (WUE, δ13C), and WUE plasticity to drought in Arabidopsis thaliana

Flowering time and water-use efficiency (WUE) are two ecological traits that are important for plant drought response. To understand the evolutionary significance of natural genetic variation in flowering time, WUE, and WUE plasticity to drought in Arabidopsis thaliana, we addressed the following questions: (1) How are ecophysiological traits genetically correlated within and between different soil moisture environments? (2) Does terminal drought select for early flowering and drought escape? (3) Is WUE plasticity to drought adaptive and/or costly? We measured a suite of ecophysiological and reproductive traits on 234 spring flowering accessions of A. thaliana grown in well-watered and season-ending soil drying treatments, and quantified patterns of genetic variation, correlation, and selection within each treatment. WUE and flowering time were consistently positively genetically correlated. WUE was correlated with WUE plasticity, but the direction changed between treatments. Selection generally favored early flowering and low WUE, with drought favoring earlier flowering significantly more than well-watered conditions. Selection for lower WUE was marginally stronger under drought. There were no net fitness costs of WUE plasticity. WUE plasticity (per se) was globally neutral, but locally favored under drought. Strong genetic correlation between WUE and flowering time may facilitate the evolution of drought escape, or constrain independent evolution of these traits. Terminal drought favored drought escape in these spring flowering accessions of A. thaliana. WUE plasticity may be favored over completely fixed development in environments with periodic drought.     

QUANTITATIVE GENETICS OF SEQUENTIAL LIFE-HISTORY AND JUVENILE TRAITS IN THE PARTIALLY SELFING PERENNIAL,AQUILEGIA CAERULEA

We determined the genetic basis of several traits related to overall fitness of Aquilegia caerulea, a perennial herb of the Rocky Mountains in western North America. To obtain measures of heritability relevant to the evolutionary potential of wild populations, we performed full and partial diallel crosses and studied progeny performance in the field. Based on a joint analysis of two designs with a total of 18 parents and 102 crosses, we detected significant maternal variance for seed mass and emergence time, but this component was negligible for later-expressed traits. Low heritability and evidence that maternal effects on seed mass are largely environmental suggest that in this population there is little evolutionary potential for change in seed mass under conditions experienced during the study. Seed mass varied depending on particular combinations of parents and cross direction. Such an interaction can have several different biological interpretations, including that particular maternal parents selectively provision embryos sired by particular pollen genotypes. Width of the first true leaf after 4 wk of growth and leaf size of juvenile plants at years one and two were significantly heritable and positively genetically correlated. Juvenile survival exhibited significant dominance variance, as expected from evidence of inbreeding depression in this trait. In contrast, for other traits that exhibit inbreeding depression in this population (seed mass and third-year leaf size), dominance variance was negligible.     

Why do evening people consider themselves more intelligent than morning individuals? The role of big five,narcissism, and objective cognitive ability

ABSTRACT

Morningness-eveningness, or chronotype, reflects the timing of sleep-wake patterns across a 24-hour day. Extant research has revealed that chronotype correlates with numerous psychological constructs including cognitive ability. In the current research, we examined how people with different chronotypes perceive their intelligence. We expected eveningness to be positively associated with subjectively assessed intelligence (SAI) because evening chronotypes demonstrate slightly higher intelligence than morning individuals. Furthermore, we considered personality traits (Big Five and narcissism) and objective intelligence (measured with standardized tests of fluid and verbal IQ) as potential variables that could account for this relationship. Across two studies (N = 504 and 232), we found that eveningness was associated with higher SAI. This relationship remained significant even after controlling for objective intelligence. In Study 1, we also found that when conscientiousness and neuroticism were analyzed together with chronotype, the magnitude of positive association between eveningness and SAI increased. Furthermore, Study 2 revealed that evening individuals exhibited higher narcissism, which fully accounted for their intelligence self-views. In the discussion, we speculate that daily struggles of evening chronotypes to function in morning-oriented society give them a basis to think positively about their intelligence to the extent of positive bias.     

Combining oxytocin administration and positive emotion inductions: Examining social perception and analytical performance

Intranasal administration of the hypothalamic neuropeptide oxytocin (OT) has, in some studies, been associated with positive effects on social perception and cognition. Similarly, positive emotion inductions can improve a range of perceptual and performance-based behaviors. In this exploratory study, we examined how OT administration and positive emotion inductions interact in their associations with social and analytical performance. Participants (N = 124) were randomly assigned to receive an intranasal spray of OT (40 IU) or placebo and then viewed one of three videos designed to engender one of the following emotion states: social warmth, pride, or an affectively neutral state. Following the emotion induction, participants completed social perception and analytical tasks. There were no significant main effects of OT condition on social perception tasks, failing to replicate prior research, or on analytical performance. Further, OT condition and positive emotion inductions did not interact with each other in their associations with social perception performance. However, OT condition and positive emotion manipulations did significantly interact in their associations with analytical performance. Specifically, combining positive emotion inductions with OT administration was associated with worse analytical performance, with the pride induction no longer benefiting performance and the warmth induction resulting in worse performance. In sum, we found little evidence for main or interactive effects of OT on social perception but preliminary evidence that OT administration may impair analytical performance when paired with positive emotion inductions.     

Evolutionary rates for multivariate traits: the role of selection and genetic variation

A fundamental question in evolutionary biology is the relative importance of selection and genetic architecture in determining evolutionary rates. Adaptive evolution can be described by the multivariate breeders'' equation (), which predicts evolutionary change for a suite of phenotypic traits () as a product of directional selection acting on them (β) and the genetic variance–covariance matrix for those traits (G). Despite being empirically challenging to estimate, there are enough published estimates of G and β to allow for synthesis of general patterns across species. We use published estimates to test the hypotheses that there are systematic differences in the rate of evolution among trait types, and that these differences are, in part, due to genetic architecture. We find some evidence that sexually selected traits exhibit faster rates of evolution compared with life-history or morphological traits. This difference does not appear to be related to stronger selection on sexually selected traits. Using numerous proposed approaches to quantifying the shape, size and structure of G, we examine how these parameters relate to one another, and how they vary among taxonomic and trait groupings. Despite considerable variation, they do not explain the observed differences in evolutionary rates.     

The Good,the Bad,and the Ugly: Agonistic Behaviour in Juvenile Crocodilians

We examined agonistic behaviour in seven species of hatchling and juvenile crocodilians held in small groups (N = 4) under similar laboratory conditions. Agonistic interactions occurred in all seven species, typically involved two individuals, were short in duration (5–15 seconds), and occurred between 1600–2200 h in open water. The nature and extent of agonistic interactions, the behaviours displayed, and the level of conspecific tolerance varied among species. Discrete postures, non-contact and contact movements are described. Three of these were species-specific: push downs by C. johnstoni; inflated tail sweeping by C. novaeguineae; and, side head striking combined with tail wagging by C. porosus. The two long-snouted species (C. johnstoni and G. gangeticus) avoided contact involving the head and often raised the head up out of the way during agonistic interactions. Several behaviours not associated with aggression are also described, including snout rubbing, raising the head up high while at rest, and the use of vocalizations. The two most aggressive species (C. porosus, C. novaeguineae) appeared to form dominance hierarchies, whereas the less aggressive species did not. Interspecific differences in agonistic behaviour may reflect evolutionary divergence associated with morphology, ecology, general life history and responses to interspecific conflict in areas where multiple species have co-existed. Understanding species-specific traits in agonistic behaviour and social tolerance has implications for the controlled raising of different species of hatchlings for conservation, management or production purposes.     

Pride and shame: Key components of a culturally universal status management system

We apply recent adaptationist theories about the emotions “pride” and “shame” to the domain of hierarchical status and test the hypothesis that pride and shame are distinct components of a culturally universal status-management system. Using an international dataset containing ratings of the status impacts of 240 personal characteristics within 14 nations (N = 2751), we found that (i) the status impacts of personal characteristics were strongly intercorrelated across nations (rs = 0.79–0.98); (ii) American's (N = 222) forecasts of the pride or shame they would experience if they exhibited those same personal characteristics closely tracked the status impacts across nations (|rs| = 0.74–0.98); and (iii) pride differentially tracked status gains, while shame differentially tracked status losses. These findings provide strong supporting evidence for the existence of a universal grammar of status criteria, and suggest that pride and shame are key components of a culturally universal status management system.     

The costs of dominance: testosterone, cortisol and intestinal parasites in wild male chimpanzees

Background

Male members of primate species that form multi-male groups typically invest considerable effort into attaining and maintaining high dominance rank. Aggressive behaviors are frequently employed to acquire and maintain dominance status, and testosterone has been considered the quintessential physiological moderator of such behaviors. Testosterone can alter both neurological and musculoskeletal functions that may potentiate pre-existing patterns of aggression. However, elevated testosterone levels impose several costs, including increased metabolic rates and immunosuppression. Cortisol also limits immune and reproductive functions.

Methods

To improve understanding of the relationships between dominance rank, hormones and infection status in nonhuman primates, we collected and analyzed 67 fecal samples from 22 wild adult male chimpanzees (Pan troglodytes schweinfurthii) at Ngogo, Kibale National Park, Uganda. Samples were analyzed for cortisol and testosterone levels as well as intestinal parasite prevalence and richness. 1,700 hours of observation data were used to determine dominance rank of each animal. We hypothesized that dominance rank would be directly associated with fecal testosterone and cortisol levels and intestinal parasite burden.

Results

Fecal testosterone (but not cortisol) levels were directly associated with dominance rank, and both testosterone and cortisol were directly associated with intestinal parasite richness (number of unique species recovered). Dominance rank was directly associated with helminth (but not protozoan) parasite richness, so that high ranking animals had higher testosterone levels and greater helminth burden.

Conclusions

One preliminary interpretation is that the antagonist pleiotropic effects of androgens and glucocorticoids place a cost on attaining and maintaining high dominance rank in this species. Because of the costs associated with elevated steroid levels, dominance status may be an honest signal of survivorship against helminth parasites.     

Caring across Boundaries versus Keeping Boundaries Intact: Links between Moral Values and Interpersonal Orientations

Prior work has established robust diversity in the extent to which different moral values are endorsed. Some people focus on values related to caring and fairness, whereas others assign additional moral weight to ingroup loyalty, respect for authority and established hierarchies, and purity concerns. Five studies explore associations between endorsement of distinct moral values and a suite of interpersonal orientations: Machiavellianism, prosocial resource distribution, Social Dominance Orientation, and reported likelihood of helping and not helping kin and close friends versus acquaintances and neighbors. We found that Machiavellianism (Studies 1, 3, 4, 5) (e.g., amorality, controlling and status-seeking behaviors) and Social Dominance Orientation (Study 4) were negatively associated with caring values, and positively associated with valuation of authority. Those higher in caring values were more likely to choose prosocial resource distributions (Studies 2, 3, 4) and to report reduced likelihood of failing to help kin/close friends or acquaintances (Study 4). Finally, greater likelihood of helping acquaintances was positively associated with all moral values tested except authority values (Study 4). The current work offers a novel approach to characterizing moral values and reveals a striking divergence between two kinds of moral values in particular: caring values and authority values. Caring values were positively linked with prosociality and negatively associated with Machiavellianism, whereas authority values were positively associated with Machiavellianism and Social Dominance Orientation.     

Preservation of winter social dominance status in Brent Geese Branta bernicla bernicla within and across winters

Dominant and subordinate individuals in a group may benefit from the stability of the social dominance organisation, avoiding excessive waste of time and energy in aggressive interactions and reducing injury risks. Nevertheless, the likely evolutionary incentive for individuals to become, and furthermore to stay, dominant may destabilise such dominance hierarchies. In this context, the relative importance of fixed (e.g. sex, morphological size) and fluctuating (e.g. body condition, mating status, reproductive success, social unit size) traits influencing the establishment and preservation of dominance relationships could play a key role in group structure. We investigated the relative role of fixed and fluctuating traits on social status in Dark-bellied Brent Geese Branta bernicla bernicla which form large fairly unstable groups both within and across winters. We compared individual dominance scores of ringed Brent Geese during four consecutive winters. Brent Geese conserved their dominance score within a given winter irrespective of their age but were generally unable to conserve it across consecutive winters. As winter dominance scores correlated best with social unit size, dominance status thus appeared to be mostly a by-product of a fluctuating trait: breeding success in the previous summer. When we considered only adults that had the same social unit size during two consecutive winters, we observed a significant preservation of dominance scores. This result suggests that a fixed trait such as sex or morphological size may still play a role in setting dominance status.     

Female macaques compete for ‘power’ and ‘commitment’ in their male partners

The formation of male-female social bonds and the resulting competition among females for male partners is a core element of human societies. While female competition for a male partner outside the mating context is well studied in humans, evidence from non-human primates is scarce, and its evolutionary roots remain to be explored. We studied two multi male – multi female groups of wild Assamese macaques (Macaca assamensis), a species where females gain benefits from selectively affiliating with particular males. Using a behavioral data set collected over several years, we tested whether females competed over access to male social partners, whether success in competition was driven by female dominance rank, and which male traits were most attractive for females. We found assortative bonding by dominance rank between females and males, which together with females initiating and maintaining contact suggests direct female competition over males. Two male traits independently predicted male attractiveness to females: (1) current dominance rank, a measure of 'power' or a male's ability to provide access to resources, and (2) prior male affiliation with immatures, a measure of a male's potential paternal proclivity or 'commitment' to infant care. Both traits have been consistently identified as drivers of female partner choice in humans. Our study adds to the evidence that female competition for valuable male partners is not unique to humans, suggesting deep evolutionary origins of women's mate choice tendencies for ‘power’ and ‘commitment’.     

Non-additive genetic variation in growth,carcass and fertility traits of beef cattle

Background

A better understanding of non-additive variance could lead to increased knowledge on the genetic control and physiology of quantitative traits, and to improved prediction of the genetic value and phenotype of individuals. Genome-wide panels of single nucleotide polymorphisms (SNPs) have been mainly used to map additive effects for quantitative traits, but they can also be used to investigate non-additive effects. We estimated dominance and epistatic effects of SNPs on various traits in beef cattle and the variance explained by dominance, and quantified the increase in accuracy of phenotype prediction by including dominance deviations in its estimation.

Methods

Genotype data (729 068 real or imputed SNPs) and phenotypes on up to 16 traits of 10 191 individuals from Bos taurus, Bos indicus and composite breeds were used. A genome-wide association study was performed by fitting the additive and dominance effects of single SNPs. The dominance variance was estimated by fitting a dominance relationship matrix constructed from the 729 068 SNPs. The accuracy of predicted phenotypic values was evaluated by best linear unbiased prediction using the additive and dominance relationship matrices. Epistatic interactions (additive × additive) were tested between each of the 28 SNPs that are known to have additive effects on multiple traits, and each of the other remaining 729 067 SNPs.

Results

The number of significant dominance effects was greater than expected by chance and most of them were in the direction that is presumed to increase fitness and in the opposite direction to inbreeding depression. Estimates of dominance variance explained by SNPs varied widely between traits, but had large standard errors. The median dominance variance across the 16 traits was equal to 5% of the phenotypic variance. Including a dominance deviation in the prediction did not significantly increase its accuracy for any of the phenotypes. The number of additive × additive epistatic effects that were statistically significant was greater than expected by chance.

Conclusions

Significant dominance and epistatic effects occur for growth, carcass and fertility traits in beef cattle but they are difficult to estimate precisely and including them in phenotype prediction does not increase its accuracy.     

Social dialect and men's voice pitch influence women's mate preferences

Low male voice pitch may communicate potential benefits for offspring in the form of heritable health and/or dominance, whereas access to resources may be indicated by correlates of socioeconomic status, such as sociolinguistic features. Here, we examine if voice pitch and social dialect influence women's perceptions of men's socioeconomic status and attractiveness. In Study 1, women perceived lower pitched male voices as higher in socioeconomic status than higher pitched male voices. In Study 2, women independently perceived lower pitched voices and higher status sociolinguistic dialects as higher in socioeconomic status and attractiveness. We also found a significant interaction wherein women preferred lower pitched men's voices more often when dialects were lower in sociolinguistic status than when they were higher in sociolinguistic status. Women also perceived lower pitched voices as higher in socioeconomic status more often when dialects were higher in sociolinguistic status than when lower in sociolinguistic status. Finally, women's own self-rated socioeconomic status was positively related to their preferences for voices with higher status sociolinguistic dialects, but not to their preferences for voice pitch. Hence, women's preferences for traits associated with potentially biologically heritable benefits, such as low voice pitch, are moderated by the presence of traits associated with resource accrual, such as social dialect markers. However, women's preferences for language markers of resource accrual may be functionally independent from preferences for potential biological indicators of heritable benefits, such as voice pitch.     

Intraspecies variation in dominance style of Macaca fuscata

Knowledge of intraspecific variation is important to test the evolutionary basis of covariation in primate social systems, yet few reports have focused on it, even in the best-studied species of the Macaca genus. We conducted a comparative study of the dominance styles among three provisioned, free-ranging groups of Japanese macaques at Shodoshima Island, Takasakiyama Mountain and Shiga Heights, and collected standard data on aggressive and affiliative behavior during a period of 5 years. Our data in the Takasakiyama and Shiga groups support previous studies showing that Japanese macaques typically have despotic social relations; nevertheless, our data in the Shodoshima group are inconsistent with the norm. The social traits of Shodoshima monkeys suggested that: (1) their dominance style is neither despotic nor tolerant but is intermediate between the two traits; (2) some measures of dominance style, e.g., frequency and duration of social interactions, covary as a set of tolerant traits in Shodoshima monkeys. This study suggests broad intraspecific variation of dominance style in Japanese macaques as can be seen in some other primate species.