Dominance relationships and patterns of aggression in a bachelor group of Sorraia horses (Equus caballus)

The influence of individual factors on dominance rank and the relationship between rank distance and patterns of aggression predicted by models of evolutionarily stable strategies (ESS) of animal conflict were investigated in a managed bachelor group of Sorraia horses, Equus caballus. The group was composed of four to six stallions 3- to 12-years-old during the study period. The dominance hierarchy was significantly linear and rank was not related to age, weight, height or aggressiveness. Frequency and intensity of agonistic interactions were low, but higher-ranking stallions did not receive lower aggressiveness than lower-ranking stallions. There was some evidence that dominance relationships were more contested among close-ranking stallions, as predicted. Agonistic-related interactions among close-ranking stallions served similar functions to those among distant-ranking stallions, but the latter interacted more frequently than expected for access to resting sites and/or resting partners. Therefore, we found some evidence that agonistic-related interactions among distant-ranking stallions play a larger role in providing access to valuable and defendable resources than those among close-ranking stallions. Nevertheless, the fact that space to escape from aggression was limited and breeding access was independent from dominance rank may have reduced the benefits relative to costs of aggression and therefore limited the occurrence of contests over dominance and resources.     

Play in adult bonobos (Pan paniscus): modality and potential meaning

The aim of this study was to thoroughly investigate social play and its modalities among adult bonobos. We evaluated how play intensity varies according to the sex-class combination of the playmates and we also performed an analysis on social locomotor-rotational movements (L-R play) and contact interactions (C play). Rough and gentle play sessions were performed with comparable frequencies by male-female and female-female adult dyads, with play signals unlikely when the playmates strongly differed in age and in rank position. L-R play rates did not differ according to the sex-combination of the players; in contrast, C play sessions were particularly frequent among females. Play faces (play signals) were significantly higher during C play than L-R play sessions, thus suggesting that playmates assess reciprocally yet safely their relationships by using facial displays to avoid any kind of misunderstanding. Play was positively correlated with grooming and contact sitting interactions, suggesting that it may be used as a social enhancer. Finally, we found no correlation between both play contexts (L-R and C play) and age, size and rank differences of the players. In conclusion, we suggest that bonobos with their egalitarian society, peculiar social structure, and playful tendency represent an attractive testing subject to examine empirically many emerging hypotheses on adult play behavior.     

Affiliative relationships among Sorraia mares: influence of age, dominance, kinship and reproductive state

The influence of individual factors on dominance rank and the relationship between rank distance and patterns of aggression predicted by models of evolutionarily stable strategies (ESS) of animal conflict were investigated in a managed bachelor group of Sorraia horses, Equus caballus. The group was composed of four to six stallions 3- to 12-years-old during the study period. The dominance hierarchy was significantly linear and rank was not related to age, weight, height or aggressiveness. Frequency and intensity of agonistic interactions were low, but higher-ranking stallions did not receive lower aggressiveness than lower-ranking stallions. There was some evidence that dominance relationships were more contested among close-ranking stallions, as predicted. Agonistic-related interactions among close-ranking stallions served similar functions to those among distant-ranking stallions, but the latter interacted more frequently than expected for access to resting sites and/or resting partners. Therefore, we found some evidence that agonistic-related interactions among distant-ranking stallions play a larger role in providing access to valuable and defendable resources than those among close-ranking stallions. Nevertheless, the fact that space to escape from aggression was limited and breeding access was independent from dominance rank may have reduced the benefits relative to costs of aggression and therefore limited the occurrence of contests over dominance and resources.     

Dominance rank predicts social network position across developmental stages in rhesus monkeys

Social network analysis is increasingly common in studying complex interactions among individuals. Across a range of primates, high-ranking adults are generally more socially connected, which results in better fitness outcomes. However, it still remains unclear whether this relationship between social network position and dominance rank emerges in infancy and whether, in species with a social transmission of dominance rank, social network positions are driven by the presence of the mother. To fill this gap, we first explored whether dominance ranks were related to social network position, measured via eigenvector centrality, in infants, juveniles, and adults in a troop of semi-free-ranging rhesus macaques (Macaca mulatta). We then examined relationships between dominance rank and eigenvector centrality in a peer-only group of yearlings who were reared with their mothers in either a rich, socially complex environment of multigenerational (MG) kin support or a unigenerational group of mothers and their infants from birth through 8 months. In Experiment 1, we found that mother's network position predicted offspring network position and that dominants across all age categories were more central in affiliative networks (social contact, social grooming, and social play). Experiment 2 showed that high-ranking yearlings in a peer-only group were more central only in the social contact network. Moreover, yearlings reared in a socially complex environment of MG kin support were more central. Our findings suggest that the relationship between dominance rank and social network position begins early in life, and that complex early social environments can promote later social competency. Our data add to the growing body of evidence that the presence/absence of the mother and kin influence how dominance rank affects social network position. These findings have important implications for the role of caregivers in the social status of developing primates, which ultimately ties to health and fitness outcomes.     

Looking like mother makes mallard ducklings dominant over their siblings

Colour variation in time and space among animals may affect social relationships such as pairing and dominance interactions. For instance, some birds are naturally sensitive to leg colour, with some colours being more visible or attractive than others. The colour of the leg-rings used to mark birds may thus be related to behavioural and reproductive variables. Most studies have investigated this effect for adults during reproduction, but leg-ring colour may also affect the behaviour of young birds. We tested the potential effect of leg-ring colours on the within-brood dominance hierarchy of mallard (Anas platyrhynchos) ducklings while each brood formed a stable and exclusive family unit with its mother. Ducklings did not acquire a within-brood dominance rank according to the colour of their own ring. This result suggests that mallards may not have a sensory bias for a given colouration. However, ducklings wearing a ring of the same colour as one of the two rings of their mother were dominant over their siblings. We discuss the potential behavioural and methodological implications of this result.     

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.     

Play behavior in immature moor macaques (Macaca maura) and Japanese macaques (Macaca fuscata)

Play is widespread across mammalian taxa, but species strongly vary in the ways they play. In less despotic primate species (i.e., with less steep dominance hierarchies, less severe conflicts, and more reconciliation), play has been described as being more frequent, cooperative, and freely expressed. To study the link between social play and dominance style, we compared play behavior in free-ranging infants, juveniles and subadults of more despotic Japanese macaques (Macaca fuscata, N = 24) and less despotic moor macaques (Macaca maura, N = 17). We found interspecific differences in play behavior that corresponded with the contrasting dominance styles of the study species, largely confirming our predictions. In particular, moor macaques spent a larger proportion of time in solitary and social play than Japanese macaques, while Japanese macaques spent a larger proportion of time in grooming interactions. In moor macaques, play sessions included more players, a larger variety of play behaviors, greater play face rates, a greater proportion of time in contact play, and a higher rate of reciprocal play-biting than in Japanese macaques. Aggressive escalations were not common, but more frequent in Japanese macaques. Finally, a higher frequency of play faces during play sessions predicted the occurrence of more reciprocal play-bites, but not the proportion of time spent in contact play behaviors. Additional studies on other groups and species will allow a better understanding of the link between dominance style and social play.     

灵长类社会玩耍的行为模式、影响因素及其功能风险

社会玩耍是指两个或两个以上个体共同参与的一种互作性玩耍行为,个体间的行为彼此适应并相互影响。社会玩耍行为在灵长类物种的社会交往过程中普遍发生,作为未成年个体一种重要的发育行为,其对个体的生存技能和成年后的繁殖成功具有重要影响。灵长类物种的社会玩耍不仅仅表现为追逐、摔跤、跳跃等一些常见行为,部分物种还发展出自己特有的行为。一般而言,社会玩耍在婴儿后期和青少年早期的发生频率最高,然后随着年龄增长直到成年时期,这类行为的平均发生频率将逐渐下降。未成年雄性个体要比同年龄段的雌性个体更喜欢玩耍,但常因物种、研究对象年龄等因素表现不同甚至相反;很多物种的个体喜欢与有亲缘关系的个体玩耍,或者与性别相同、年龄相仿、等级相近的个体玩耍。总之,非人灵长类个体社会玩耍的发育不但受环境参量如食物、场地等的影响,而且还与个体的年龄、性别、等级、亲缘关系等社群因素紧密相关。未成年个体在玩耍过程中,获得了身体机能的快速发育、完善了生存技能、建立了个体间的友好关系、增强了认识自身及适应周围环境的能力,从而为顺利过渡到成年期和履行自己的职能打好基础,但有时却需要承担玩耍过程中受伤甚至死亡的风险。玩耍作为灵长类社会的一种行为文化,对其研究有助于对人类自身行为进化的不断认识,相信这方面的理论将会得到后来者的不断创新和丰富,也期望这方面的理念及经验能被及时运用到保护繁育等实践活动中。     

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 relationships in a herd of Sorraia horses Part I. Correlates of social dominance and contexts of aggression

Factors related to dominance rank and the functions of aggression were studied in a herd of Sorraia horses, Equus caballus, under extensive management. Subjects were 10 adult mares 5-18 years old and a stallion introduced into the group for breeding. Dominance relationships among mares were clear, irrespective of rank difference, and remained stable after introduction of the stallion. The dominance hierarchy was significantly linear and rank was positively correlated with age and total aggressiveness. Higher-ranking mares received lower frequency and intensity of agonistic interactions. Nevertheless, higher-ranking dominants were not more likely to elicit submission from their subordinates than lower-ranking dominants. Neither close-ranking mares nor mares with less clear dominance relationships were more aggressive towards each other. Agonistic interactions seemed to be used more importantly in regulation of space than to obtain access to food or to reassert dominance relationships. Contexts of aggression were related to mare rank. The results suggest that dominance relationships based on age as a conventional criterion were established to reduce aggressiveness in a herd where the costs of aggression are likely to outweigh the benefits.     

Sex differences in the impact of social status on hair cortisol concentrations in rhesus monkeys (Macaca mulatta)

Social status impacts stress in primates, but the direction of the effect differs depending on species, social style, and group stability. This complicates our ability to identify broadly applicable principles for understanding how social status impacts health and fitness. One reason for this is the fact that social status is often measured as linear dominance rank, yet social status is more complex than simply high or low rank. Additionally, most research on social status and health ignores the effects of sex and sex-specific relationships, despite known differences in disease risk, coping strategies, and opposite-sex dominance interactions between males and females in many species. We examine the influence of social status, sex, and opposite-sex interactions on hair cortisol concentrations (HCC) in a well-studied species, rhesus macaques, where the literature predicts low ranking individuals would experience more chronic stress. Animals in three captive, seminaturalistic social groups (N = 252; 71 male) were observed for 6 weeks to obtain metrics of social status (rank and dominance certainty [DC]). DC is a measure of one's fit within the hierarchy. Hair samples were collected from each subject and analyzed for HCC. Generalized linear mixed models were used to examine (a) whether rank, DC, or sex predicted HCC; (b) whether same- or opposite-sex dominance relationships differentially impacted HCC; and (c) whether aggressive interactions initiated or received could explain any observed relationships. Results indicated that DC, not rank, predicted HCC in a sex-specific manner. For males, high HCC were predicted by receiving aggression from or having high DC with other males as well as having low DC with females. For females, only high DC with males predicted high HCC. These results likely relate to sex-specific life history pattern differences in inherited versus earned rank that are tied to female philopatry and male immigration.     

Is the adipose fin and the lower jaw (kype) related to social dominance in male Arctic charr Salvelinus alpinus?

This study reports on the importance of the size of the kype (lower jaw) and the adipose fin for establishing and maintaining social dominance in pair-wise interactions among size-matched, reproductively active male Arctic charr Salvelinus alpinus. The size of these traits seems not to have a large influence on establishing dominance, but after 4 days of social interactions, and after dominance rank is established, subordinate males show reduced size of their adipose fins and kypes relative to that of dominant males. Consequently, these traits seem to be costly labile characters that could be of importance in inter and intra-sexual evaluations of individual quality.     

Method for Assigning Categorical Rank in Female <Emphasis Type="Italic">Pan troglodytes schweinfurthii</Emphasis> via the Frequency of Approaches

Establishing the order of a dominance hierarchy among female chimpanzees (Pan troglodytes) is complicated by the fact that they often forage solitarily, and aggressive interactions between them occur infrequently. Authors of previous studies have typically ranked females via the direction of submissive pant-grunts and the outcome of agonistic interactions. Given that higher rank correlates with higher reproductive success in female chimpanzees, assessing rank is important but may be limited by sparsely populated dominance matrices. I tested the hypothesis that rank predicts the direction of female approaches. There is a significant relationship among Gombe females between the frequency with which a female was approached and her dominance rank. Dominant females approached other females less often than they were approached. Though approached frequencies failed to meet the criteria for formal rank indicators, they may be useful as real indicators of subordination. Because approach interactions occur far more frequently than pant-grunts, they may be useful in assigning categorical rank when traditional dominance metrics are limited.     

Adolescent male chimpanzees do not form a dominance hierarchy with their peers

Dominance hierarchies are a prominent feature of the lives of many primate species. These hierarchies have important fitness consequences, as high rank is often positively correlated with reproduction. Although adult male chimpanzees strive for status to gain fitness benefits, the development of dominance relationships is not well understood. While two prior studies found that adolescent males do not display dominance relationships with peers, additional research at Ngogo in Kibale National Park, Uganda, indicates that adolescents there form a linear dominance hierarchy. These conflicting findings could reflect different patterns of rank acquisition across sites. An alternate possibility arises from a recent re-evaluation of age estimates at Ngogo and suggests that the report describing decided dominance relationships between adolescent males may have been due to the accidental inclusion of young adult males in the sample. To investigate these issues, we conducted a study of 23 adolescent male chimpanzees of known age during 12 months at Ngogo. Adolescent male chimpanzees exchanged pant grunts, a formal signal of submission, only 21 times. Recipients of pant grunts were late adolescent males, ranging between 14 and 16 years old. In contrast, younger adolescent males never received pant grunts from other males. Aggression between adolescent males was also rare. Analysis of pant grunts and aggressive interactions did not produce a linear dominance hierarchy among adolescent males. These data indicate that adolescent male chimpanzees do not form decided dominance relationships with their peers and are consistent with the hypothesis that the hierarchy described previously at Ngogo resulted from inaccurate age estimates of male chimpanzees. Because dominance relationships develop before adulthood in other primates, our finding that adolescent male chimpanzees do not do so is surprising. We offer possible explanations for why this is the case and suggest future studies that may help clarify the matter.     

Patterns of coalition formation by adult female baboons in Amboseli, Kenya

Coalitionary support in agonistic interactions is generally thought to be costly to the actor and beneficial to the recipient. Explanations for such cooperative interactions usually invoke kin selection, reciprocal altruism or mutualism. We evaluated the role of these factors and individual benefits in shaping the pattern of coalitionary activity among adult female savannah baboons, Papio cynocephalus, in Amboseli, Kenya. There is a broad consensus that, when ecological conditions favour collective defence of resources, selection favours investment in social relationships with those likely to provide coalitionary support. The primary features of social organization in female-bonded groups, including female philopatry, linear dominance hierarchies, acquisition of maternal rank and well-differentiated female relationships, are thought to be functionally linked to the existence of alliances between females. Female savannah baboons display these characteristics, but the frequency and function of their coalitionary aggression is disputed. In our five study groups, 4-6% of all disputes between females led to intervention by third parties. Adult females selectively supported close maternal kin. There was no evidence that females traded grooming for support or reciprocated support with nonkin. High-ranking females participated in coalitionary aggression most frequently, perhaps because they derived more benefits from group membership than other females did or could provide support at lower cost. Females typically supported the higher ranking of two contestants when they intervened in disputes between subordinates, so most coalitions reinforced the existing dominance hierarchy. Results indicate that female baboons participate in coalitionary aggression in a manner strongly influenced by nepotism and individual benefits.     

Dominance relationships among one-male units in a provisioned free-ranging band of the Sichuan snub-nosed monkeys (Rhinopithecus roxellana) in the Qinling Mountains, China

We studied the dominance relationships among one-male units (OMUs) in a provisioned free-ranging band of the Sichuan snub-nosed monkeys (Rhinopithecus roxellana) in the Qinling Mountains of central China from 2001 to 2005. The band was composed of 6-8 OMUs that stayed in the band for several years. Linear dominance orders could be detected using displacement interactions with directional asymmetry among OMUs in 82.3+/-5% of interactions, and ambiguous and reversed interactions in 17.7+/-5%. The dominance rank of OMUs was positively related with the duration of their stay in the band, and this may be attributed to the association of the resident male with adult females, rather than the fighting ability of resident males, as males do not fight seriously with each other. Subordinate units were observed to merge with dominant units resulting in an elevation of their rank order. The linear dominance relationship among OMUs in the Sichuan snub-nosed monkeys may have evolved as a result of competition for preferred food trees.     

Acquisition of dominance status affects maze learning in mice

Learning is likely to be costly and thus subject to trade-off with other components of life history. An obvious prediction, therefore, is that investment in learning, and thus learning performance, will vary with individual life history strategy and the reproductive value of the learning outcome. We tested this idea in the context of social dominance in male laboratory mice, using a simple radial maze paradigm to compare the ability of high- and low-ranking male mice to track changing food location. We tested animals in randomly selected pairs before and after establishing aggressive rank relationships to distinguish intrinsic differences in learning ability from those attributable to acquiring high or low rank. There was no difference in learning between later dominants and subordinates prior to establishing rank relationships. After pairing, however, dominants showed a significantly greater percentage of correct responses, with the difference being greatest earlier in a sequence of trials. The percentage of correct responses also increased with the amount of aggression initiated during pairing. The results thus appeared to reflect a state-dependent change in learning associated with the aggressive social relationships formed during pairing.     

Worker dominance and policing in the ant Temnothorax unifasciatus

In many species of eusocial Hymenoptera, conflict about the production of males is resolved through “policing.” Recent studies in wasps and the ant Temnothorax unifasciatus suggest that in these species policing workers are dominant themselves and selfishly increase their own chances of later becoming fertile. Policing may therefore to some extent be associated with dominance and selfishness, and dominance and policing behaviour are indeed difficult to distinguish and often not mutually exclusive. Moreover, selfish policing requires that workers form rank orders already in the presence of the queen. Here, we try to allocate aggressiveness by workers towards policing and/or dominance behaviour and investigate whether hierarchies based on subtle, non-aggressive interactions exist in queenright colonies of the ant T. unifasciatus. We either split colonies into a queenright and queenless halve or temporarily removed the queen from complete colonies, which in both cases allows a few dominant workers to lay eggs in the queenless colony. Reunification of colony halves and return of the queen to orphaned colonies led to aggression against those workers that had become fertile during the absence of the queen. Dominant workers in reunited, split colonies were more severely attacked than those in orphaned colonies after return of the queen. Furthermore, we observed that workers, which later became dominant egg layers under queenless conditions, have more contact with the queen than other workers. Both results corroborate the existence of rank relationships among workers in queenright colonies and show that results from policing experiments may be affected by the disturbance of pre-existing hierarchies through colony splitting.     

Does Male Badge Size Signal Status in Small Island Populations of House Sparrows,Passer domesticus?

The relationship between dominance and throat badge size, body size, and body condition was examined in three small island populations of house sparrows, Passer domesticus, on the coast of northern Norway. Individual dominance rank in males was determined by observation of agonistic interactions in an artificial observation cage (one replicate in Dec. and one in Mar. in two populations and one replicate in Mar. in the third population). Previous studies had indicated that badge size signals status in house sparrows. In this study, because of the small population sizes and stability in their membership, we expected that prior information about fighting ability would reduce the importance of badge size for the outcome of conflicts in favour of traits most directly associated with fighting ability. In two of five replicates, significant relationships were found between components of body size and dominance rank, but in opposite directions. The relationship between badge size and dominance rank was consistently positive, but non-significant. In a pooled sample of the three independent replicates, badge size was the best predictor of dominance, and no other variable explained any variance in dominance. However, the coefficient of determination was low and linearity of dominance hierarchies was poor. We therefore suggest that individual recognition may influence the dominance relationships.     

Strong Competition Does Not Always Predict Play Asymmetry: The Case of South American Sea Lions (Otaria flavescens)

Although play fighting has been studied for over a century in both human and non‐human animals, quantitative data on marine mammals are still scarce. Here, we investigated play fighting in South American sea lions (Otaria flavescens), one of the most sexually dimorphic species with an extreme polygynous mating system, high levels of both intra‐ and inter‐sexual competition. All these features make South American sea lions a good model species to test some predictions on play fighting. Our results indicate play is restricted to juveniles, being inhibited among adults, and as to be expected in a species that shows a high degree of sexual dimorphism, it is mainly expressed in males. Even though playful interactions were punctuated by competitive behaviours, animals played in a highly symmetric way and were able to adjust their competitive playful interactions in a flexible manner and so reduce the risk of escalation to a minimum level. They were highly selective in their choice of playmates by limiting the number of players per session and playing more with age‐matched companions and friends. All these factors taken together are probably at the basis of the low risk of escalation recorded during the study. This result is predictive of a high ability and motivation of these animals to engage in play behaviour which can have a possible role not only in the acquisition of dominance status, but also in establishing and maintaining social relationships, an unexpected role in a so highly competitive species.