Dominance,aggression and physiological stress in wild male Assamese macaques (Macaca assamensis)

In group-living animals relative rank positions are often associated with differences in glucocorticoid output. During phases of social stability, when dominance positions are clear and unchallenged, subordinates often face higher costs in terms of social stress than dominant individuals. In this study we test this prediction and examine additional potential correlates of stress, such as reproductive season, age and amount of aggression received in wild, seasonally breeding Assamese macaques (Macaca assamensis). During a mating and a non-mating season we collected 394 h of focal observational data and 440 fecal samples of six adult and six large subadult males living in a multimale–multifemale group in their natural habitat in northeastern Thailand. The mating season was characterized by a general increase in aggressive behavior and glucocorticoid excretion across all males compared to the non-mating season. Among adult males, mating season glucocorticoid levels were significantly negatively related with dominance rank and positively with the amount of aggression received. Both relationships were non-significant among large subadult males. Thus, our results suggest that in adult Assamese macaques a high dominance position is not associated with high costs. Low costs of dominance might be induced by strong social bonds among top-ranking males, which exchange frequent affiliative interactions and serve as allies in coalitionary aggression against potentially rank-challenging subordinate males.     

Advanced age influences chimpanzee behavior in small social groups

Management strategies for captive chimpanzees (Pan troglodytes) must begin to take into account the increasing age of the captive chimpanzee population. This study represents a baseline assessment of the relationship between advancing age and behavior among male and female chimpanzees living in pairs and trios in indoor/outdoor runs. Data collected on 14 old individuals (30–44 years old) and 20 younger adult individuals (11–22 years old) totaled 240 hours. Levels of agonistic and affiliative social behavior, non‐social activity, abnormal behaviors, and behavioral indicators of anxiety were evaluated. In the same captive setting, the behavior of old chimpanzees was significantly different from younger chimpanzees. Old chimpanzees showed less aggression and moved about their enclosures less. Old females behaved submissively more often than younger adult females; the reverse was found among males. However, affiliative social behavior occurred at similar levels in old and younger adult chimpanzees, implying continued need for social housing with advancing age. The effect of enrichment devices may differ for aged female chimpanzees, given their submissiveness and the lower levels of object manipulation found in aged subjects. These results suggest that aging in chimpanzees may be accompanied by altered patterns of social interaction, requiring careful attention to the compatibility of social partners. Zoo Biol 19:111–119, 2000. © 2000 Wiley‐Liss, Inc.     

Child social ethology and peer relations: a developmental review of methodology and findings

During the last three decades, developmental research has increasingly emphasized the relevance of peer relations in children’s socialization. However, most studies of child development still focus upon individual differences in social status, tacitly neglecting relational constraints inherent in the ecology of the peer group. In contrast, socioethological approaches have stressed that natural groups provide a variety of distinct social roles that may have a differential impact upon individual growth and development. However, ethological analyses have often been limited to aggressive relations and group dominance structures. Comparable studies of affiliative organization have been hampered by the paucity of models for the study of cohesive social structure, only recently having begun to overcome this obstacle. A potential third dimension of preschool peer group ethology, object use, has been relatively neglected in studies of peer relations. However, recent research suggests that object use is a salient and important component of the social world of the young child. This article reviews methodology and recent findings in the area of peer relations and discusses the developmental implications of this work. Received: 8 December 1998 / Received in revised form: 26 March 1999 / Accepted: 31 March 1999     

Psychosocial and hormonal aspects of hierarchy formation in groups of male squirrel monkeys

Accounts of primate social dominance hierarchies often imply that the achievement of superior status is a “goal”, akin to a valued resource or commodity, and that hierarchies emerge in multimale groups from prolonged competitive conflicts over social status. This possibility is not consistent with our observations of five newly formed triads of adult male squirrel monkeys (Saimiri sciureus). Stable linear hierarchies based on clear asymmetries in the direction of intermale agonism and genital displays were established quickly, with virtually no reciprocal fighting, and in the absence of rank-related differences in plasma cortisol or testosterone. Although affiliative social overtures were initiated more often by high-ranking and middle-ranking males, affiliative overtures were directed equally often toward all members of each group. From the outset of the study all males, regardless of rank, spent an average of 33% of their time huddling in affiliative contact with male cagemates. These results suggest that in newly formed groups of adult male squirrel monkeys, social hierarchies reflect an expedient convention that reduces conflict and facilitates the formation of small cohesive groups. © 1994 Wiley-Liss, Inc.     

Intermale affiliative behavior in ringtailed lemurs (Lemur catta) at the Beza-Mahafaly reserve, Madagascar

Intermale affiliative behavior was studied in three groups of naturally occurring ringtailed lemurs over a one-year period. The adult males in the sample exhibited affiliative behavior with all other males in their social groups; but affiliative behavior between some male-male dyads occurred markedly more often than between others. These dyads are called “preferred partnerships.” The formation of preferred partnerships did not appear to be influenced by male dominance rank or age-class. Furthermore, these partnerships were of a short-term nature, and none persisted throughout the entire study period. The following factors may explain the absence of rank or age effects, and the brevity of preferred partnerships: (1) frequent fluctuation in the male dominance hierarchy in each study group; (2) the fact that higher-ranking males may not offer specific benefits to lower-ranking males; and (3) the fact that male dispersal affected the male membership of each group over the 12-month study period. The focal males engaged in significantly more affiliative behavior during the lactation period compared with the other reproductive seasons. Migration partners may offer each other predator protection during the transfer process, and although males that transfer together do not engage in alliance or coalition behavior towards resident males, males transferring in pairs or threesomes are in a better position to spot and defend themselves against attack by resident males. Migration partners also provide social contact for each other while in the process of immigration.     

The Organization of Collective Group Movements in Wild Barbary Macaques (Macaca sylvanus): Social Structure Drives Processes of Group Coordination in Macaques

Social animals have to coordinate activities and collective movements to benefit from the advantages of group living. Animals in large groups maintain cohesion by self-organization processes whereas in smaller groups consensus decisions can be reached. Where consensus decisions are relevant leadership may emerge. Variation in the organization of collective movements has been linked to variation in female social tolerance among macaque species ranging from despotic to egalitarian. Here we investigated the processes underlying group movements in a wild macaque species characterized by a degree of social tolerance intermediate to previously studied congeneric species. We focused on processes before, during and after the departure of the first individual. To this end, we observed one group of wild Barbary macaques (Macaca sylvanus) in the Middle Atlas, Morocco using all-occurrence behaviour sampling of 199 collective movements. We found that initiators of a collective movement usually chose the direction in which more individuals displayed pre-departure behavior. Dominant individuals contributed to group movements more than subordinates, especially juveniles, measured as frequencies of successful initiations and pre-departure behaviour. Joining was determined by affiliative relationships and the number of individuals that already joined the movement (mimetism). Thus, in our study group partially shared consensus decisions mediated by selective mimetism seemed to be prevalent, overall supporting the suggestion that a species’ social style affects the organization of group movements. As only the most tolerant species show equally shared consensus decisions whereas in others the decision is partially shared with a bias to dominant individuals the type of consensus decisions seems to follow a stepwise relation. Joining order may also follow a stepwise, however opposite, relationship, because dominance only determined joining in highly despotic, but not in intermediate and tolerant species.     

Aggression and social support predict long‐term cortisol levels in captive tufted capuchin monkeys (Cebus [Sapajus] apella)

Many nonhuman primates live in complex social groups in which they regularly encounter both social stressors such as aggression and social support such as that provided by long‐term affiliative relationships. Repeated exposure to social stressors may result in chronically elevated cortisol levels that can have deleterious physical effects such as impaired immune function, cardiovascular disease, and reduced brain function. In contrast, affiliative social relationships may act as a buffer, dampening the release of cortisol in response to acute and chronic stressors. Understanding how social stressors and social support predict cortisol levels is therefore essential to understanding how social situations relate to health and welfare. We studied this relationship in 16 socially housed captive brown capuchin monkeys (Cebus [Sapajus] apella) by comparing long‐term hair cortisol levels with behavioral measures of social stress (dominance rank, rank certainty, and amount of aggression received) and social support (amount of affiliation and centrality in the affiliative social network of the group). Dominance rank, rank certainty, amount of affiliation, and age were not significant predictors of long‐term cortisol levels in this population. Instead, long‐term cortisol levels were positively related to the amount of aggression received and negatively related to centrality in the affiliative social network of the group. This pattern may be attributed to the species’ socially tolerant dominance system and to the availability of social support across the dominance hierarchy.     

Social organization of a stable natal group of captive Guyanese squirrel monkeys (<Emphasis Type="Italic">Saimiri sciureus sciureus</Emphasis>)

Socioecological models suggest competition for food, foraging efficiency, predation, infanticide risk, and the costs of dispersal regulate primate social structure and organization. Wild populations of squirrel monkeys (Saimiri spp.) appear to conform to the predictions of the predation/competition socioecological model (Sterck et al. in Behav Ecol Sociobiol 41:291–309, 1997) and the dispersal/foraging efficiency model (Isbell in Kinship and behavior in primates. Oxford University, New York, pp 71–108, 2004). However, squirrel monkeys in captivity are reported to maintain patterns of social behavior observed in their wild conspecifics despite different food distribution, predation risk, and dispersal options. This behavioral similarity suggests squirrel monkeys’ social behavior has limited flexibility to respond to environmental changes. In this study, we experimentally evaluated the flexibility of social behavior within a captive group of S. sciureus. First, we determined whether dominance and affiliative relationships observed under normal laboratory conditions (with abundant, widely distributed, food; no dispersal option; and no predators) better matched published reports of relationships among wild conspecifics or the predictions of the predation/competition model. Second, we made preferred food items defensible to determine whether dominance interactions would become more frequent and linear, as predicted by the model. The model correctly predicted rates of dominance behavior in both conditions and a linear hierarchy in the defensible food condition but did not predict the consistent affiliative relationships and linear dominance hierarchy observed in normal lab conditions. Although hierarchies were linear and male dominant, manipulating food distribution changed the dominant individual within each sex. Our findings suggest interaction rates adapt more rapidly than social structure to environmental changes in Saimiri and recommend caution in interpreting tests of socioecological models.     

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.     

Social behavior in fission–fusion groups of red uakari monkeys (Cacajao calvus ucayalii)

Primates living in large groups that divide to forage must have social systems compatible with this mode of living. Uakari monkeys (Cacajao spp.) live in large groups and exhibit a form of fission–fusion grouping, but their social organization is poorly understood. We present some of the first data on social behavior for this genus based on a study on Cacajao calvus ucayalii. They traveled in multimale multifemale groups of highly variable sizes, with bachelor units on the periphery. Adult males were affiliative, and adult females associated with more than one adult male. Adult females typically traveled with their dependent offspring and an older juvenile within the group. In parties of two or more males, individuals engaged in previously unreported display behaviors and acted together to aggressively chase other males. Breeding was seasonal, and mating occurred away from other group members. We speculate on the social organization of C. calvus ucayalii, in which dispersal may be bisexual and peripheral males are affiliative with one another. Affiliated males appear to cooperate in fighting and displaying to other males for access to females during the breeding season. Am. J. Primatol. 71:976–987, 2009. © 2009 Wiley‐Liss, Inc.     

Female social dominance in two Eulemur species with different social organizations

Female social dominance is rare in mammals, but common in lemurs. We investigated social dominance in two Eulemur species; the polygynous crowned lemur (E. coronatus) and the monogamous red‐bellied lemur (E. rubriventer), using four and two social groups, respectively. We collected data on agonistic interactions and two types of affiliative behavior (grooming and maintaining spatial proximity). We used a combination of focal watches of individuals, instantaneous scan‐sampling of groups, and all‐occurrence of some behaviors in groups. We found that overall rates of agonistic interactions were higher in E. coronatus, and they also had more decided intersexual agonistic interactions than E. rubriventer. However, in both species the females won the vast majority of these agonistic interactions. E. coronatus females were groomed more often by males than vice versa, whereas no sex differences in grooming were observed in E. rubriventer. We found that males were responsible for maintaining spatial proximity in E. coronatus whereas in E. rubriventer, females were responsible. In one group of E. coronatus, the male was overweight and dominant to the female and this is the first observation of male dominance in a lemur species typically described as female dominant. We suggest that body weights in captivity be monitored for maintaining normal dominance relationships. Overall, agonistic behaviors were consistent with clear female social dominance in both E. coronatus and E. rubriventer. The affiliative behaviors also provided clear evidence for female dominance E. coronatus, but not for E. rubriventer. Zoo Biol 0: 1–14, 2007. © 2007 Wiley‐Liss, Inc.     

Importance of cooperation and affiliation in the evolution of primate sociality

The idea that competition and aggression are central to an understanding of the origins of group‐living and sociality among human and nonhuman primates is the dominant theory in primatology today. Using this paradigm, researchers have focused their attention on competitive and aggressive behaviors, and have tended to overlook the importance of cooperative and affiliative behaviors. However, cooperative and affiliative behaviors are considerably more common than agonistic behaviors in all primate species. The current paradigm often fails to explain the context, function, and social tactics underlying affiliative and agonistic behavior. Here, we present data on a basic question of primate sociality: how much time do diurnal, group‐living primates spend in social behavior, and how much of this time is affiliative and agonistic? These data are derived from a survey of 81 studies, including 28 genera and 60 species. We find that group‐living prosimians, New World monkeys, Old World monkeys, and apes usually devote less than 10% of their activity budget to active social interactions. Further, rates of agonistic behaviors are extremely low, normally less than 1% of the activity budget. If the cost to the actors of affiliative behavior is low even if the rewards are low or extremely variable, we should expect affiliation and cooperation to be frequent. This is especially true under conditions in which individuals benefit from the collective environment of living in stable social groups. Am J Phys Anthropol 128:84–97, 2005. © 2005 Wiley‐Liss, Inc.     

Yellow-tailed woolly monkey (Lagothrix flavicauda) proximal spacing and forest strata use in La Esperanza,Peru

Inter-individual spacing of primates and their use of forest strata depend on temporal and spatial changes in the environment and on predator avoidance, group demography, and social conditions. Greater proximity also increases the chances of agonistic and affinitive contact between individuals. I collected behavioral data for three groups of yellow-tailed woolly monkeys (Lagothrix flavicauda) by instantaneous sampling of focal animals for 15 months in La Esperanza, northeastern Peru. By use of combined data for all groups I examined the effects of season, activity, and age/sex class on nearest-neighbor distances and forest strata use. Small differences were observed for nearest-neighbor distances, forest strata use by different age/sex classes, and activity. Adult males had the lowest contact index scores. Contact index scores were low for juvenile females, for which nearest-neighbor distances were largest. Very little aggressive behavior was observed. Focal animals preferred upper levels of the forest with little difference in height for different activities. Lagothrix flavicauda have very cohesive groups with little seasonal or activity-dependent difference between nearest-neighbor distances or proximity. These results suggest that this species has less variable social organization and greater group cohesion than other Atelini. However, more studies are needed on other populations of L. flavicauda to better determine the species’ social organization. Studies are also required to determine the extent to which dispersal times and kinship affect proximity, nearest-neighbor distances, and aggression.     

Evaluating Dominance Styles in Assamese and Rhesus Macaques

Researchers have suggested that several types of agonistic and affiliative behavior covary as a set of species-specific traits, and have used the term dominance style to describe the covariation. We compared measures of dominance style between a group of Assamese macaques (Macaca assamensis) and a group of rhesus macaques (M. mulatta), though kinship information was unknown. Assamese and rhesus female-female dyads each showed a low proportion of counter aggression and a low conciliatory tendency, suggesting that they have despotic social relationships. They also showed a despotic pattern on several other types of agonistic and affiliative behavior, such as approach outcomes and grooming distributions, which is consistent with the covariation of dominance style traits. Assamese male-male dyads showed relatively high levels of reconciliation and counter aggression versus other macaque males portrayed in the literature, suggesting that Assamese males have a tolerant dominance style. Insofar as macaque dominance style depends on the behavior of females, we suggest that Assamese macaques, like rhesus macaques, have despotic social relationships, which contrasts with evidence of a strong correlation between phylogeny and dominance style in macaques. Further, our results indicate that strong male bonding and tolerant dominance relationships among males are independent of female dominance style. Lastly, some measures of agonistic behavior, such as rate of aggression or proportion of bites, are likely altered in competitive environments and thus are not useful indicators of dominance style.     

Female social relationships in a captive group of Campbell's monkeys (Cercopithecus campbelli campbelli)

A study group of Campbell's monkeys (Cercopithecus c. campbelli) provided data on affiliative and agonistic relationships between females. Over a period of two years (involving 111 hr), we conducted observations of a captive group which had a composition similar to wild groups. We were able to identify a monitor-adjust social system with frequent affiliative interactions, directed gazing and avoidances rather than aggressive acts. We described long-term differentiated affiliative bonds: adult females interacting more often with some group mates than others, especially if they were relatives. Interactions between matrilines concerned essentially play and young adult females. We found a significant linear hierarchy of dominance with rare reversals and a stable intermatriline dominance. In contrast to other single-male groups, our adult male was socially integrated into the group although this may have been because of the housing conditions. Comparisons with the social organization of other captive and wild guenon groups are discussed.     

Dominance, aggression and physiological stress in wild male Assamese macaques (Macaca assamensis)

In group-living animals relative rank positions are often associated with differences in glucocorticoid output. During phases of social stability, when dominance positions are clear and unchallenged, subordinates often face higher costs in terms of social stress than dominant individuals. In this study we test this prediction and examine additional potential correlates of stress, such as reproductive season, age and amount of aggression received in wild, seasonally breeding Assamese macaques (Macaca assamensis). During a mating and a non-mating season we collected 394 h of focal observational data and 440 fecal samples of six adult and six large subadult males living in a multimale–multifemale group in their natural habitat in northeastern Thailand. The mating season was characterized by a general increase in aggressive behavior and glucocorticoid excretion across all males compared to the non-mating season. Among adult males, mating season glucocorticoid levels were significantly negatively related with dominance rank and positively with the amount of aggression received. Both relationships were non-significant among large subadult males. Thus, our results suggest that in adult Assamese macaques a high dominance position is not associated with high costs. Low costs of dominance might be induced by strong social bonds among top-ranking males, which exchange frequent affiliative interactions and serve as allies in coalitionary aggression against potentially rank-challenging subordinate males.     

Children's understanding of dominance and prestige in China and the UK

Individuals can gain high social rank through dominance (based on coercion and fear) and prestige (based on merit and admiration). We conducted a cross-cultural developmental study and tested 5- to 12-year-olds, and adults in the UK and China, aiming to determine (a) the age at which children distinguish dominance and prestige, and (b) the influence of cultural values on rank-related reasoning. We specifically tested participants in China because of the value of prestigious individuals modestly yielding to subordinates, a social skill that becomes more salient with age. In both populations, the distinction between dominance and prestige emerged at five years, and improved over childhood. When reasoning about a resource conflict between a high-ranking party and a subordinate, adults in both countries expected high-rank individuals to win, although Chinese adults were less likely to do so regarding prestigious individuals. Across the two countries, younger children (5–7 years) responded similarly to each other, not favoring either party as the winner. Older children (9–12 years), however, diverged. Those in the UK chose the high-rank party, while those in China made no systematic inference. Overall, our findings suggest that while children distinguish prestige and dominance comparably in the two countries, they develop culturally-influenced expectations about the behavior of high-rank individuals.     

Social Structure Affects Initiations of Group Movements but Not Recruitment Success in Japanese Macaques (<Emphasis Type="Italic">Macaca fuscata</Emphasis>)

Research on collective movements has often focused on the sociodemographic parameters explaining the success of some individuals as leaders or initiators of collective movements. Several of these studies have shown the influence of social structure, through kinship and affiliative relationships, on the organization of collective movements. However, these studies have been conducted on semi-free-ranging groups of macaques that were not faced with a natural environment and its constraints. In the socially intolerant rhesus macaque (Macaca mulatta) the success of an initiator correlates with its hierarchical rank, the most dominant individuals being the most successful. We investigated the collective movements of another socially intolerant macaque species, Japanese macaques, in the wild, to assess whether the social structure was still a determinant factor under natural conditions. In line with previous studies of macaques, we found that social structure drove the organization of collective movements. More dominant individuals initiated more collective movements. However, dominance did not affect the success of an initiation, i.e., the number of individuals joining. In addition, kinship strongly constrained the associations observed in females during collective movements. These results reflect the social structure of Japanese macaques, in which strong power asymmetry and kinship relationships constrain the majority of interactions between individuals within the group. Moreover, these results are similar to those observed in semi-free-ranging rhesus macaques and support the hypothesis of an effect of social determinants on collective movements of primates even under natural conditions.     

Paternal Relatedness Predicts the Strength of Social Bonds among Female Rhesus Macaques

Forming strong, equitable, and enduring social bonds with a few individuals in a group carries adaptive benefits in terms of increased longevity, offspring survival and paternity success in birds and mammals, including humans. These recent insights generated a new interest in the factors creating variation in the strength of social relationships. Whether and how animals discriminate paternal kin from non-kin and bias their social behavior accordingly is being debated. This study explores the relative importance of dominance rank, age, maternal and paternal relatedness in shaping dyadic affiliative relationships in a group of 30 captive rhesus macaque females. The strength of social relationships, measured by the composite sociality index from observational data, was independently predicted in GLMMs by both maternal and paternal relatedness as well as rank similarity. In addition, social bonds between paternal half-sisters were stronger than between distantly related kin suggesting that females biased their affiliative effort towards paternal relatives. Despite identical relatedness coefficients bonds between mothers and their daughters were three times as strong as those between full sisters. Together these results add to the growing body of evidence for paternal kin biases in affiliative behavior and highlight that females prefer descendent over lateral kin.     

Social behaviour of cattle in tropical silvopastoral and monoculture systems

Silvopastoral systems can be a good alternative for sustainable livestock production because they can provide ecosystem services and improve animal welfare. Most farm animals live in groups and the social organization and interactions between individuals have an impact on their welfare. Therefore, the objective of this study was to describe and compare the social behaviour of cattle (Bos indicus×Bos taurus) in a silvopastoral system based on a high density of leucaena (Leucaena leucocephala) combined with guinea grass (Megathyrsus maximus), star grass (Cynodon nlemfuensis) and some trees; with a monoculture system with C. nlemfuensis, in the region of Merida, Yucatán. Eight heifers in each system were observed from 0730 to 1530 h each day for 12 consecutive days during the dry season and 12 consecutive days during the rainy season. The animals followed a rotation between three paddocks, remaining 4 days in each paddock. The vegetation was characterized in the paddocks of the silvopastoral system to estimate the average percentage of shade provided. To make a comparison between systems, we used a t test with group dispersion, and Mann–Whitney tests with the frequency of affiliative and agonistic behaviours. We assessed differences in linearity and stability of dominance hierarchies using Landau’s index and Dietz R-test, respectively. The distance of cows with respect to the centroid of the group was shorter, and non-agonistic behaviours were 62% more frequent in the intensive silvopastoral system than in the monoculture one. Heifers in the silvopastoral system had a more linear and non-random dominance hierarchy in both seasons (dry season: h’=0.964; rainy season: h’=0.988), than heifers in the monoculture system (dry season: h’=0.571, rainy season: h’=0.536). The dominance hierarchy in the silvopastoral system was more stable between seasons (R-test=0.779) than in the monoculture system (R-test=0.224). Our results provide the first evidence that heifers in the silvopastoral system maintain more stable social hierarchies and express more sociopositive behaviours, suggesting that animal welfare was enhanced.