Testosterone and Linear Social Dominance Status in Captive Male Dabbling Ducks in Winter

Dominance hierarchies play an important role in avoidance and/or solving conflicts in gregarious species. In dabbling ducks (Anas species), dominance allows for feeding‐site monopolization in winter quarters where resources are generally limited. In addition, male social rank should theoretically favour access to mates. Dominance rank can be associated with morphological traits, and is often correlated with aggressiveness, a behavioural trait generally related to high testosterone levels. In this study, we investigated the existence of a winter group structure based on dominance relationships and tested for a linear hierarchy, in three species of captive male dabbling ducks (mallard Anas platyrhynchos, pintail A. acuta and wigeon A. penelope). We then analysed the relationship between dominance ranks, morphological parameters and testosterone levels measured in early (Oct.) and mid‐winter (Dec./Jan.). We found that the three male groups of the three species exhibited a linear hierarchy. Testosterone levels differed during winter and between species. Morphologic measurements, body mass and body condition were not correlated with individual dominance ranks, whereas dominant males had higher testosterone levels than subordinates. The slopes of the relationships were similar between species and winter period, but the y‐intercepts differed between species and between early and mid‐winter phases. The linear hierarchy found in the three species indicates that dominance relationships strongly structure dabbling duck groups in winter. Lack of correlation between rank and morphological characters, but correlation of rank with testosterone levels suggests that social rank is more dependent on behavioural traits such as aggressive behaviour. The differences between species and winter periods are discussed in relation to migration and wintering phenology.     

Social correlates of testosterone and ornamentation in male mandrills

We investigated relationships between fecal androgen concentrations, facial coloration and behaviour in semi-free-ranging male mandrills. We found that fecal androgen levels were significantly positively related to dominance rank, independent of rank stability and the mating period, suggesting that male mandrills live in a permanently aggressive context in which they must actively maintain their dominance status. Facial red coloration was also significantly related to both fecal androgen levels and rank, with high ranking males having both higher androgen levels and redder faces, although dominant males did not always have the highest androgen levels or the reddest faces. Predictive relationships between androgen levels, coloration and rank were short-term. Androgen concentrations and facial redness both increased in the presence of receptive females, as did the former during periods of rank instability. We conclude that male facial redness is likely to represent an honest signal (to other males) of current androgen status, competitive ability and willingness to engage in fights and that females may also use this to assess male condition. Further, our findings provide support for the “challenge hypothesis” as originally proposed for birds by Wingfield.     

Social and environmental factors affecting fecal glucocorticoids in wild, female white-faced capuchins (Cebus capucinus)

Assessing glucocorticoid levels in free-ranging nonhuman primates provides a means to determine the social and environmental stress load for individuals. We investigated the effect of four proximate variables--reproductive state, season, male rank stability, and dominance rank--on the level of fecal glucocorticoids (cortisol metabolites) in eight adult female white-faced capuchin monkeys in Costa Rica. Reproductive state, season, and male rank stability significantly affected fecal glucocorticoids while female dominance rank did not. Cortisol levels were significantly higher in pregnant females as compared with lactating or other reproductive states. Cortisol levels were higher among females during the dry season compared with the wet season, suggesting a metabolic adaptation to maintain homeostasis in drier, hotter conditions. Although unfamiliar males present a greater infanticidal threat than do familiar ones, we found that females experienced higher glucocorticoid levels during male rank instability events, regardless of whether the alpha male role was taken over by a familiar or an unfamiliar male. Our findings provide important benchmark and comparative data for future studies on the variables that affect glucocorticoid levels in this species and other mammals.     

Anogenital distance as a measure of male competitive ability in Rwenzori Angolan colobus

Anogenital distance (AGD) is positively correlated to fetal androgen exposure and developmental masculinization in mammals. Independent of overall body size, AGD shows a strong positive correlation with male fertility and in rodents, AGD is a good indicator of male competitive ability and is associated with female choice. We hypothesized that AGD will also predict male competitive ability in non-human primates. To test this, we measured AGD noninvasively with a parallel laser in a wild population of Angolan colobus monkeys (Colobus angolensis ruwenzorii) in Uganda and correlated to it to their social structure. C. angolensis ruwenzorii form a multilevel society with both one-male/multifemale units (OMUs) and multimale/multifemale units (MMUs). We compared AGD in males from five OMUs and six MMUs and related it to their fecal androgen metabolite concentrations, dominance rank and body size, and to the number of females in their unit. Males in OMUs had greater access to females, so were predicted to have longer AGDs, but this was not found. AGD also did not correlate overall with mean fecal androgen metabolites in MMUs. However, AGD was correlated with dominance rank in MMUs, demonstrating that higher-ranking males in these multimale units had longer AGDs. Body size did not show the same relationship with dominance rank, suggesting that male rank was not just a reflection of absolute male size. Our findings indicate that AGD predicts male competitive ability in this species and that it may be a useful correlate throughout the non-human primates. These results also support the idea that prenatal androgen exposure increases the likelihood of the expression of behaviors that maintain high dominance rank.     

Social dominance does not increase oxidative stress in a female dominance hierarchy of an African cichlid fish

In many group living animal species, individuals use aggression to gain and maintain social dominance to secure access to ecological resources and potential mates. While social dominance has many fitness benefits, there are also potential costs associated with frequent agonistic interactions and status display. One potential cost of social dominance is oxidative stress, the imbalance of reactive oxygen species and antioxidant capacity. In the cichlid species Astatotilapia burtoni, dominant males are aggressive, hold a breeding territory, and have an activated reproductive system resulting in larger gonads. Subordinate males are submissive, school with females, and are nonreproductive. Females are submissive under natural conditions, but in a female-only group, a dominance hierarchy will form with dominant females taking on male-typical behaviours including aggression, territory defence, and increased androgen levels. However, in contrast to males, social dominance is not linked to increased activation of the reproductive system in females, allowing us to test whether social dominance alone exposes individuals to increased oxidative stress. We compared dominant and subordinate females in female-only groups in five markers of oxidative stress. Dominant females did not have higher levels of oxidative damage compared to same-sex subordinates. This result contrasted to the trend in males in which dominant males had higher oxidative damage than their subordinate counterparts. Our findings suggest that the oxidative cost of social dominance is limited and support the notion that previously reported associations between high rank and increased oxidative stress is most likely driven by increased investment in reproduction.     

Beneficial effect of hot spring bathing on stress levels in Japanese macaques

The ability of animals to survive dramatic climates depends on their physiology, morphology and behaviour, but is often influenced by the configuration of their habitat. Along with autonomic responses, thermoregulatory behaviours, including postural adjustments, social aggregation, and use of trees for shelter, help individuals maintain homeostasis across climate variations. Japanese macaques (Macaca fuscata) are the world’s most northerly species of nonhuman primates and have adapted to extremely cold environments. Given that thermoregulatory stress can increase glucocorticoid concentrations in primates, we hypothesized that by using an available hot spring, Japanese macaques could gain protection against weather-induced cold stress during winter. We studied 12 adult female Japanese macaques living in Jigokudani Monkey Park, Japan, during the spring birth season (April to June) and winter mating season (October to December). We collected faecal samples for determination of faecal glucocorticoid (fGC) metabolite concentrations by enzyme immunoassay, as well as behavioural data to determine time spent in the hot springs, dominance rank, aggression rates, and affiliative behaviours. We used nonparametric statistics to examine seasonal changes in hot spring bathing, and the relationship between rank and air temperature on hot spring bathing. We used general linear mixed-effect models to examine factors impacting hormone concentrations. We found that Japanese macaques use hot spring bathing for thermoregulation during the winter. In the studied troop, the single hot spring is a restricted resource favoured by dominant females. High social rank had both costs and benefits: dominant females sustained high fGC levels, which were associated with high aggression rates in winter, but benefited by priority of access to the hot spring, which was associated with low fGC concentrations and therefore might help reduce energy expenditure and subsequent body heat loss. This unique habit of hot spring bathing by Japanese macaques illustrates how behavioural flexibility can help counter cold climate stress, with likely implications for reproduction and survival.     

Within-brood social status and consequences for winter hierarchies amongst Mallard Anas platyrhynchos ducklings

Juveniles of many birds establish dominance hierarchies within family social units, only to leave and compete to acquire dominance status in new social groups. Little is known about the role of sex, body mass, size or experience during the duckling period on subsequent dominance rank and adult social relationships. We used captive Mallard Anas platyrhynchos ducklings to test for the role of individual characteristics and growth parameters in establishing within-brood hierarchies, the maintenance of within-brood hierarchies in the subsequent wintering group and differences in social ranks between broods. Strong stable linear hierarchies were present within each brood and, later, within each phase of the winter. There was a reorganisation of the hierarchical order between the duckling period and early winter, but only few modifications afterwards during the winter. None of the tested “hatching”, “duckling” and “adult” traits explained either the within-brood or the winter hierarchies, but winter rank was related to brood of origin with ducklings from the same brood having similar social ranks. These differences between broods were maintained through the whole winter in most cases, though one brood drastically progressed in the hierarchy during late-winter. These results suggest that the factors affecting the establishment of social relationships within broods differ from those in winter groups, and that brood-related mechanisms influence social relationships during winter. We discuss our results in the light of direct and indirect maternal influence.     

Stress, social behaviour, and secondary sexual traits in a male primate

We examined variation in glucocorticoid levels in the mandrill, a brightly coloured primate species, to identify major social influences on stress hormones, and investigate relationships among glucocorticoid levels, testosterone and secondary sexual ornamentation. We collected a total of 317 fecal samples for 16 adult male mandrills over 13 months, including mating and non-mating periods and periods of both dominance rank stability and instability, and compared fecal glucocorticoid levels with dominance rank, rank stability, presence of receptive females, gastro-intestinal parasite infection, fecal testosterone and facial red coloration. Glucocorticoid levels did not vary systematically with dominance rank, but increased when the dominance hierarchy was unstable, and increased in the presence of receptive females. The relationship between dominance rank and glucocorticoid levels changed direction according to the stability of the dominance hierarchy: glucocorticoid levels were higher in subordinate males under stable conditions, but under conditions of instability higher ranking males had higher glucocorticoid levels. The influence of dominance rank also interacted with the presence of receptive females: glucocorticoids were higher in dominant males than in subordinates, but only during mating periods, suggesting that dominant males are more stressed than subordinates during such periods. These findings support previous studies showing that the relationship between glucocorticoids and dominance rank in male baboons is dependent on the social environment. We also found that males with higher glucocorticoids suffered a higher diversity of gastrointestinal parasite infection, in line with evidence that glucocorticoids suppress the immune system in other species. However, we found no support for the stress-mediated immunocompetence handicap hypothesis for the evolution of condition-dependent ornaments: glucocorticoid and testosterone levels were positively related, rather than the negative relationship predicted by the hypothesis, and we found no relationship between red colour and glucocorticoid levels, suggesting that glucocorticoids do not play a role in translating social conditions or physical health into ornament expression in this species.     

Females competing to reproduce: dominance matters but testosterone may not

The associations among aggression, testosterone (T), and reproductive success have been well studied, particularly in male birds. In many species, males challenged with simulated or real territorial intrusions increase T and levels of aggression, outcomes linked to higher dominance status and greater reproductive success. For females, the patterns are less clear. Females behave aggressively towards one another, and in some species, females respond to a social challenge with increases in T, but in other species they do not. Prior work on female dark-eyed juncos (Junco hyemalis) had shown that experimental elevation of T increases social status and intrasexual aggression. Here, we conducted two experiments designed to answer three questions: Are endogenous concentrations of T associated with dominance status in captive female juncos? Does dominance status influence readiness to breed in female juncos? And do captive females increase T in response to a challenge? In the first experiment, we introduced two females to a breeding aviary, allowed them to form a dominance relationship and then introduced a male. We found that dominant females were more likely to breed than subordinates, but that dominance status was not predicted by circulating T. In the second experiment, we allowed a resident male and female to establish ownership of a breeding aviary (territory) then introduced a second, intruder female. We found that resident females were aggressive towards and dominant over intruders, but T did not increase during aggressive interactions. We suggest that during the breeding season, intrasexual aggression between females may influence reproductive success, but not be dependent upon fluctuations in T. Selection may have favored independence of aggression from T because high concentrations of T could interfere with normal ovulation or produce detrimental maternal effects.     

Influence of reproductive season and rank on fecal glucocorticoid levels in free-ranging male Verreaux's sifakas (Propithecus verreauxi)

Studies in anthropoid primates and other mammals suggest that reproductive season, rank, reproductive skew, aggression received, and social support are the major factors influencing glucocorticoid output. In which way these are also affecting adrenal function in lemurid primates has been studied rarely. Here, we examine the influence of reproductive season and rank on glucocorticoid output in male sifakas (Propithecus verreauxi), a species characterized by high breeding seasonality, a hierarchy among males and extreme reproductive skew towards dominant males. We established a fecal assay for non-invasively monitoring adrenal activity and collected 315 fecal samples during the reproductive and birth season from 10 male sifakas living in 5 groups in Western Madagascar. We found a significant effect of season on glucocorticoid output, with males exhibiting higher fecal glucocorticoid levels during the reproductive compared to the birth season in conjunction with an increase in overall aggression rates during the former period. Moreover, our data indicate a significant effect of rank on adrenocortical activity with dominant males exhibiting higher glucocorticoid levels than subordinate males in the reproductive season. However, dominant males did not differ significantly in rates of initiated or received aggression and rates of affiliative behavior from subordinates but showed significantly lower rates of submission. Given their highly formalized dominance relationships, we conclude that higher glucocorticoid output in dominant males during the 4-month reproductive season is likely related to higher energetic demands necessary to cope with the challenges of male reproduction rather than to physical demands of increased fighting frequency to maintain dominance status. High rank in sifakas may thus carry high costs, which, however, may be outweighed by monopolization of almost all paternities. In sum, our data generally support the findings on the relationship between environmental and social factors and glucocorticoid output found in non-lemurid primates.     

Factors affecting fecal glucocorticoid levels in semi-free-ranging female mandrills (Mandrillus sphinx)

Subordinate female cercopithecine primates often experience decreased reproductive success in comparison with high-ranking females, with a later age at sexual maturity and first reproduction and/or longer interbirth intervals. One explanation that has traditionally been advanced to explain this is high levels of chronic social stress in subordinates, resulting from agonistic and aggressive interactions and leading to higher basal levels of glucocorticoids. We assessed the relationships among fecal cortisol levels and reproductive condition, dominance rank, degree of social support, and fertility in female mandrills (Mandrillus sphinx) living in a semi-free-ranging colony in Franceville, Gabon. Lower-ranking females in this colony have a reproductive disadvantage relative to higher-ranking females, and we were interested in determining whether this relationship between dominance rank and reproductive success is mediated through stress hormones. We analyzed 340 fecal samples from 19 females, collected over a 14-month period. We found that pregnant females experienced higher fecal cortisol levels than cycling or lactating females. This is similar to results for other primate species and is likely owing to increased metabolic demands and interactions between the hypothalamus-pituitary-adrenal axis, estrogen, and placental production of corticotrophin releasing hormones during pregnancy. There was no influence of dominance rank on fecal cortisol levels, suggesting that subordinate females do not suffer chronic stress. This may be because female mandrills have a stable social hierarchy, with low levels of aggression and high social support. However, we found no relationship between matriline size, as a measure of social support, and fecal cortisol levels. Subordinates may be able to avoid aggression from dominants in the large enclosure or may react only transiently to specific aggressive events, rather than continuously expecting them. Finally, we found no relationship between fecal cortisol levels and fertility. There was no difference in fecal cortisol levels between conceptive and nonconceptive cycles, and no significant relationship between fecal cortisol level and either the length of postpartum amenorrhea or the number of cycles before conception. This suggests that the influence of dominance rank on female reproductive success in this population is not mediated through chronic stress in subordinate females, and that alternative explanations of the relationship between social rank and reproduction should be sought.     

Social rank,feeding and winter weight loss in red deer: any evidence of interference competition?

During winter at northern latitudes, large herbivores often exploit patches of concentrated, relatively high quality forage, which may lead to interference competition. The factors affecting success in contests and subsequent dominance rank, such as age and body weight, remain key issues in ungulate behavioural ecology. Maternal effects on offspring body weight are well known, but few studies have investigated if mothers social rank influence offspring rank. Moreover, no study has related dominance rank in ungulates to weight loss during winter. Outcomes of social interactions (n=7,609), feeding time and spatial position in red deer (Cervus elaphus) hinds and calves, and weight loss of calves, were registered from 1981 to 1996 at six winter-feeding sites within the county of Sør-Trøndelag in Norway. The level of aggressiveness was higher among calves than among adult hinds, and the factors determining the outcome of contests also differed. The initiator won the majority of interactions (more than 90% in both hinds and calves). Social rank was related to both age and body weight in adult hinds, and related to body weight and mother rank in calves. The relationship between feeding time and rank was non-linear. Feeding time was correlated with rank only among high ranked hinds, while there was no such relationship among low ranked hinds or calves. There was no correlation between winter weight loss and social rank in calves. Our study therefore underlines that, although frequent aggression is observed at artificial feeding sites of northern herbivores, this is not necessarily sufficient to give rise to interference competition.     

Behavioral and physiological plasticity: Rapid changes during social ascent in an African cichlid fish

In many vertebrates, reproduction is regulated by social interactions in which dominant males control access to females and food. Subordinate males that displace dominant individuals must rapidly adopt behavioral and physiological traits of the higher rank to gain reproductive success. To understand the process of phenotypic plasticity during social ascent, we analyzed the temporal expression pattern of dominance behaviors and circulating androgen levels when socially-suppressed males of an African cichlid fish Astatotilapia burtoni ascended in status. These experiments tested a prediction of the ‘challenge hypothesis’ that, during periods of social instability, male androgen levels are higher than during socially stable times. We found that socially and reproductively suppressed males perform territorial and reproductive behaviors within minutes of an opportunity to ascend in status, and that animals switch from initial expression of territorial behaviors to more reproductive behaviors during territory establishment. Following this rapid response, social stability may be achieved within 1-3 days of social ascent. Consistent with predictions of the ‘challenge hypothesis’, circulating 11-ketotestosterone (11-KT) levels were elevated within 30 min following social opportunity, coincident with increased aggressive behavior. However, territorial behaviors and serum 11-KT levels were then dissociated by 72 h after social ascent, suggesting either rapid social stability and/or increased physiological potential for androgen production. This behavioral and physiological plasticity in male A. burtoni suggests that perception of social opportunity triggers a suite of quick changes to facilitate rapid transition towards reproductive success, and reveals important features of social ascent not previously recognized.     

Habitat preference in the dark-eyed junco (Junco hyemalis): The role of photoperiod and dominance

Wild-caught dark-eyed juncos (Junco hyemalis), simultaneously presented with projected slides of their winter and summer habitats, demonstrated distinct preferences (in terms of time spent in front of particular slides) consistent with their season of capture and prior laboratory experience. Juncos caught in and kept on winter or summer photoperiods preferred winter and summer habitat slides respectively. A gradual increase in photoperiod up to natural summer conditions induced a switch from winter to summer habitat preference: a decrease in photoperiod induced a range of slide preferences correlated with the junco's size, a measure of the bird's dominance rank.     

Androgen levels and female social dominance in Lemur catta

Morphological and behavioural traits which improve agonistic power are subject to intrasexual selection and, at the proximate level, are influenced by circulating androgens. Because intrasexual selection in mammals is more intense among males, they typically dominate females. Female social dominance is therefore unexpected and, indeed, rare. Ring-tailed lemurs (Lemur catta) are sexually monomorphic primates in which all adult females dominate all males. The goal of our study was to test the prediction that female dominance in this species is associated with high androgen levels. Using two captive groups, we collected data on agonistic behaviour and non-invasively assessed their androgen concentrations in faeces and saliva by enzyme immunoassay. We found that adult female L. catta do not have higher androgen levels than males. However, during the mating season there was a twofold increase in both the androgen levels and conflict rates among females. This seasonal increase in their androgen levels was probably not due to a general increase in ovarian hormone production because those females showing the strongest signs of follicular development tended to have low androgen concentrations. At the individual level neither the individual aggression rates nor the proportion of same-sexed individuals dominated were correlated with their androgen levels. We conclude that female dominance in ring-tailed lemurs is neither based on physical superiority nor on high androgen levels and that it is equally important to study male subordination and prenatal brain priming effects for a complete understanding of this phenomenon.     

Sex-reversed correlation between stress levels and dominance rank in a captive non-breeder flock of crows

Group living has both benefits and costs to individuals; benefits include efficient acquisition of resources, and costs include stress from social conflicts among group members. Such social challenges result in hierarchical dominance ranking among group members as a solution to avoid escalating conflict that causes different levels of basal stress between individuals at different ranks. Stress-associated glucocorticoid (corticosterone in rodents and birds; CORT) levels are known to correlate with dominance rank in diverse taxa and to covary with various social factors, such as sex and dominance maintenance styles. Although there is much evidence for sex differences in the basal levels of CORT in various species, the correlation of sex differences in basal CORT with dominance rank is poorly understood. We investigated the correlation between CORT metabolites (CM) in the droppings and social factors, including rank and sex, in a captive non-breeder group of crows. In this group, all the single males dominated all the single females, and dominance ranks were stable among single males but relatively unstable among single females. CM levels and rank were significantly correlated in a sex-reversed fashion: males at higher rank (i.e., more dominant) had higher CM, whereas females at higher rank exhibited lower CM. This is the first evidence of sex-reversed patterns of CM–rank correlation in birds. The results suggest that different mechanisms of stress–dominance relationships operate on the sexes in non-breeder crow aggregations; in males, stress is associated with the cost of aggressive displays, whereas females experience subordination stress due to males' overt aggression.     

Nocturnal loss of body reserves reveals high survival risk for subordinate great tits wintering at extremely low ambient temperatures

Winter acclimatization in birds is a complex of several strategies based on metabolic adjustment accompanied by long-term management of resources such as fattening. However, wintering birds often maintain fat reserves below their physiological capacity, suggesting a cost involved with excessive levels of reserves. We studied body reserves of roosting great tits in relation to their dominance status under two contrasting temperature regimes to see whether individuals are capable of optimizing their survival strategies under extreme environmental conditions. We predicted less pronounced loss of body mass and body condition and lower rates of overnight mortality in dominant great tits at both mild and extremely low ambient temperatures, when ambient temperature dropped down to ?43 °C. The results showed that dominant great tits consistently maintained lower reserve levels than subordinates regardless of ambient temperature. However, dominants responded to the rising risk of starvation under low temperatures by increasing their body reserves, whereas subdominant birds decreased reserve levels in harsh conditions. Yet, their losses of body mass and body reserves were always lower than in subordinate birds. None of the dominant great tits were found dead, while five young females and one adult female were found dead in nest boxes during cold spells when ambient temperatures dropped down to ?43 °C. The dead great tits lost up to 23.83 % of their evening body mass during cold nights while surviving individuals lost on average 12.78 % of their evening body mass. Our results show that fattening strategies of great tits reflect an adaptive role of winter fattening which is sensitive to changes in ambient temperatures and differs among individuals of different social ranks.     

Social modulation of androgen levels in male teleost fish

Androgens are classically thought of as the sex steroids controlling male reproduction. However, in recent years evidence has accumulated showing that androgens can also be affected by the interactions between conspecifics, suggesting reciprocal interactions between androgens and behaviour. These results have been interpreted as an adaptation for individuals to adjust their agonistic motivation and to cope with changes in their social environment. Thus, male-male interactions would stimulate the production of androgens, and the levels of androgens would be a function of the stability of its social environment ['challenge hypothesis', Gen. Comp. Endocrinol. 56 (1984) 417]. Here the available data on social modulation of androgen levels in male teleosts are reviewed and some predictions of the challenge hypothesis are addressed using teleosts as a study model. We investigate the causal link between social status, territoriality and elevated androgen levels and the available evidence suggests that the social environment indeed modulates the endocrine axis of teleosts. The association between higher androgen levels and social rank emerges mainly in periods of social instability. As reported in the avian literature, in teleosts the trade-off between androgens and parental care is indicated by the fact that during the parental phase breeding males decreased their androgen levels. A comparison of androgen responsiveness between teleost species with different mating and parenting systems also reveals that parenting explains the variation observed in androgen responsiveness to a higher degree than the mating strategy. Finally, the adaptive value of social modulation of androgens and some of its evolutionary consequences are discussed.     

Status signalling, metabolic rate and body mass in the siskin: the cost of being a subordinate

The higher metabolic rate of dominant individuals, found in different species, has been interpreted as the cost that prevents subordinates from cheating by adopting large badges of status. However, an alternative prediction for status-signalling species, in which subordinates may recognize dominants, is that subordinates have the higher metabolic rate because of the greater stress of locating and actively avoiding aggressive interactions with them. In this study, the size of the black bib of the siskin, Carduelis spinus, which is a badge of dominance, was negatively correlated with metabolic rate in daylight, even when controlling for the bird's activity level in the respirometer chamber and its body mass. The size of the black bib, however, was not correlated with metabolic rate in darkness. This suggests that the difference between dominance classes is not related to intrinsic physiological differences, but that subordinates are more susceptible to stressful conditions. When controlling for metabolic rate, a positive correlation appeared between dominance status and body mass. This stresses the importance of knowing the effects of social status on energy requirements for understanding the relationship between body mass and dominance. We conclude that maintaining a high social status may be more stressful to subordinates than to dominant birds. Copyright 2000 The Association for the Study of Animal Behaviour.     

Social parameters and urinary testosterone level in male chimpanzees (Pan troglodytes)

Studies investigating relationships between social parameters (such as dominance rank, rates of aggressive and sexual behaviors) and androgen (particularly, testosterone) levels in male primates have yielded inconsistent results. In the present study, we address the relationship between androgens, male dominance rank and rank-associated behaviors in two groups of captive chimpanzees, a species characterized by a pronounced dominance hierarchy between adult males. By combining behavioral observations with urinary testosterone (T) measurements, we found that the differences in T concentrations between males were small and not obviously related to their dominance rank. T levels were not related to the rates of initiated aggression and copulatory behavior, but a significant negative relationship between male T level and the rates of strong aggression received was apparent. Our findings, combined with those of others, suggest that any relationship between dominance rank and T depends upon the extent to which individual rank-associated behaviors (e.g. aggressive/sexual) are themselves related to T.