Alpha male chimpanzee grooming patterns: implications for dominance “style”

In social primates, individuals use various tactics to compete for dominance rank. Grooming, displays and contact aggression are common components of a male chimpanzee's dominance repertoire. The optimal combination of these behaviors is likely to differ among males with individuals exhibiting a dominance “style” that reflects their tendency to use cooperative and/or agonistic dominance tactics. Here, we examine the grooming behavior of three alpha male chimpanzees at Gombe National Park, Tanzania. We found that (1) these males differed significantly in their tendency to groom with other males; (2) each male's grooming patterns remained consistent before, during and after his tenure as alpha, and (3) the three males tended to groom with high‐ middle‐ and low‐ranking partners equally. We suggest that body mass may be one possible determinant of differences in grooming behavior. The largest male exhibited the lowest overall grooming rates, whereas the smallest male spent the most time grooming others. This is probably because large males are more effective at physically intimidating subordinates. To achieve alpha status, a small male may need to compensate for reduced size by investing more time and energy in grooming, thereby ensuring coalitionary support from others. Rates of contact aggression and charging displays conformed to this prediction, suggesting that each male exhibited a different dominance “style.” Am. J. Primatol. 71:136–144, 2009. © 2008 Wiley‐Liss, Inc.     

Harassment of adults by immatures in bonobos (<Emphasis Type="Italic">Pan paniscus</Emphasis>): testing the Exploratory Aggression and Rank Improvement hypotheses

The immatures of many primate species frequently pester adult group members with aggressive behaviors referred to as a type of harassment. Although these behaviors are characteristic of immatures as they develop from infancy through adolescence, there have been few studies that specifically address the adaptive significance of harassment. Two functional hypotheses have been generated from observations of the behavior in chimpanzees. The Exploratory Aggression hypothesis describes harassment as a mechanism used by immatures to learn about the parameters of aggression and dominance behavior and to acquire information about novel, complex, or unpredictable relationships. The Rank Improvement hypothesis describes harassment as a mechanism of dominance acquisition used by immatures to outrank adults. This study investigated harassment of adults by immatures in a group of bonobos housed at the Columbus Zoo and compared the results to the predictions outlined by the Exploratory Aggression and Rank Improvement hypotheses. Although all immature bonobos in this group harassed adults, adolescents performed the behavior more frequently than did infants or juveniles and low-ranking adults were targeted more frequently than high-ranking. Targets responded more with agonistic behaviors than with neutral behaviors and the amount of harassment an individual received was significantly correlated with the amount of agonistic responses given. Furthermore, bouts of harassment were found to continue significantly more frequently when responses were agonistic than when they were neutral. Adolescents elicited mostly agonistic responses from targets whereas infants and juveniles received mostly neutral responses. These results support predictions from each hypothesis where harassment functions both as a mechanism of social exploration and as a tool to establish dominance rank.     

雄性大仓鼠体重对其斗殴行为及社会等级的作用

在实验室条件下测定雄性大仓鼠体重对社会等级和斗殴行为的作用模式,检验体重对雄性大仓鼠社会等级及斗殴行为序列具有重要影响的假设。本实验以16只成年雄性大仓鼠为目标个体,采用等级内部的线性概率、组内循环三元组数量（d）和优势等级的线性度（K）,排列个体的社会等级序位。研究结果表明,雄性大仓鼠可形成近似线性的优势等级,体重与个体的优势等级,攻击行为和胁腺标记行为均呈显著的正相关关系,与防御行为和攻击潜伏期存在显著的负相关关系。说明独居性物种大仓鼠雄体间可形成优势等级关系,体重对此关系具有重要的作用。     

Spatial clumping of food and social dominance affect interference competition among ruddy turnstones

In studying the success of foraging animals, studies of interferencecompetition have put emphasis on effects of competitor density,whereas studies of resource defense have focused on the effectsof the spatial distribution of food within patches. Very fewstudies have looked at both factors simultaneously, that is,determined whether the effects of competitor density on foragingsuccess depend on the spatial distribution of food. We studiedthe behavior and the foraging success of ruddy turnstones (Arenariainterpres) using an experiment in which we varied both the presenceof a competitor and the food distribution. Because turnstonesmay differ strongly in their relative dominance status, we alsoexperimentally varied the foragers' relative dominance status.We found that the presence of a competitor only reduced theforaging success of subordinate birds foraging at the clumpedfood distribution. At this condition, dominant and subordinatebirds differed markedly in their foraging success. Contraryto our expectations, we did not observe more agonistic behaviorat the clumped food distribution. This indicates that the amountof agonistic behavior observed may be a bad indicator of interferenceeffects. These findings have specific implications for modelsof interference competition. Most notably they show that theeffects of competitor density on agonistic behavior and foragingsuccess may well depend on the spatial distribution of foodand the foragers' relative dominance status. Additionally, ourresults suggest that social dominance will not be fully understoodwithout considering long-term processes such as the formationand maintenance of social dominance hierarchies.     

Dominance and its Behavioral Measures in a Captive Group of Bonobos (Pan paniscus)

We investigated the existence of a social dominance hierarchy in the captive group of six adult bonobos at the Planckendael Zoo. We quantified the pattern of dyadic exchange of a number of behaviors to examine to what extent each behavior fits a linear rank order model. Following de Waal (1989), we distinguish three types of dominance: agonistic dominance, competitive ability and formal dominance. Fleeing upon aggression is a good measure of agonistic dominance. The agonistic dominance hierarchy in the study group shows significant and strong linearity. The rank order was: 1. female (22 yr), 2. female (15 yr)., 3. male (23 yr.), 4. female (15 yr.), 5. male (9 yr.), 6. male (10 yr.). As in the wild, the females occupy high ranks. There is prominent but nonexclusive female agonistic dominance. Teeth-baring does not fulfil the criteria of a formal submission signal. Peering is a request for tolerance of proximity. Since its direction within dyads is consistent with that of fleeing interactions, it is a useful additional measure to determine agonistic ranks in bonobos. In competitive situations, the females acquire more food than other group members do. The rank obtained from access to food resources differs from the agonistic rank due to female intrasexual social tolerance, expressed in food sharing. We typify the dominance styles in the group as female intrasexual tolerance and male challenging of rank differences. The agonistic rank order correlates significantly with age and has a strong predictive value for other social behaviors.     

Children's verbal and nonverbal communication in a conflict situation

Same-sexed pairs of children (N = 144) were videotaped during brief play sessions involving a limited but desirable resource. Pairs varied in sex, age (preschool, kindergarten, second grade), friendship (friends vs nonfriends), and perceived dominance (dominant vs. subordinate status within the pair). Videotape coding focused on verbal and nonverbal communication during negotiations for access to the resource. Results showed that dominant children played longer with the desirable object. Negotiation strategies varied with interactional role (e.g., attempting to gain vs. retain access to object) as well as age, sex, and social relationship. The politeness of children's language varied with their affective state as indexed by use of facial expressions.     

Agonistic behavior and dominance relationships in female Phayre's leaf monkeys -- preliminary results

Socioecological theory suggests a link between the strength of competition for food/safety, rates of agonism, structure of dominance hierarchies, and dispersal among group-living females. This study presents preliminary data on agonistic behavior and dominance relationships for female Phayre's leaf monkeys (Trachypithecus phayrei), a species in which females routinely disperse. Behavioral observations were conducted on two groups (four adult females, and five adult females plus two juvenile females, respectively) at Phu Khieo Wildlife Sanctuary, northeast Thailand. Rates of agonistic behavior were analyzed from focal continuous recordings, while dominance hierarchies were constructed from all agonistic behaviors (focal and ad libitum sampling). Overall, female-female agonistic behaviors (aggression, submission, and displacements) occurred at a rate of < 0.25 interactions per hour. Agonistic interactions involving food occurred more frequently than expected based on feeding time. Females in both groups exhibited linear dominance hierarchies with some reversals, and possibly an age-inversed hierarchical structure in the larger group. The results fit well with previous results for colobine monkeys regarding frequency of interactions, displacements predominating agonistic behavior, and the possibility of an age-inversed hierarchy. The results contradict the suggested link between linearity of hierarchies and female philopatry. Future studies should consider the notion that female dispersal may coexist with linear dominance hierarchies.     

Agonistic behavior of the crayfish Euastacus armatus and Cherax destructor

The intra- and interspecific agonistic behavior of Euastacus armatus and Cherax destructor from northeastern Victoria were examined. While the agonistic patterns of E. armatus appeared similar to those shown by most crayfish, individuals of C. destructor execute an unusual, highly stylized cheliped “punch” behavior during strong interactions, along with the other behaviors seen in many species. Juvenile C. destructor exhibited gregarious behavior, tending to co-occupy burrows and being physically near each other. Tests showed that the white chelipeds which are characteristic of mature E. armatus affect the outcome of aggressive interactions. When individuals had their chelipeds whitened, they won agonistic interactions more often. This result held for both intraspecific pairings and size-matched pairings with individuals of C. destructor. Individuals of C. destructor won the majority of size-matched pairings with non-whitened individuals of E. armatus.     

Affiliative relationships and reciprocity among adult male bonnet macaques (Macaca radiata) at Arunachala Hill, India

In captivity, male bonnet macaques (Macaca radiata) frequently express "friendship" toward one another, including affiliative behavior such as huddling, grooming, coalitionary support, and sitting in close proximity. The purpose of this study was to determine whether wild adult male bonnet macaques also express "friendship" by investigating whether or not (1) adult male bonnet macaques have affiliative social relationships with other males, (2) the strength of social relationships varies among dyads, (3) there is time-matched reciprocity in allogrooming among dyads, and if so, whether the level of reciprocity occurs within a bout of grooming, a day, or over 2 months (the limit of this study), and (4) a correlation exists between the strength of social relationships and dominance ranks among adult males. Focal samples totaling 150 hr on all seven adult males in one study group were conducted to record both agonistic and affiliative interactions. Agonistic interactions were used to construct a dominance hierarchy, whereas affiliative interactions (sitting in proximity to within 1 m with and without grooming) were used to quantify the existence and strength of social bonds within dyads. Results show that adult male bonnet macaques had differentiated affiliative relationships with other males in their group. There was little reciprocity of grooming within a bout of grooming or within a day, but greater reciprocity over the study period of 2 months. There was no correlation between dominance ranking distance and the strength of affiliative relationship within dyads; however, within dyads lower-ranking males groomed higher-ranking males more than vice versa. This study suggests that friendships in male bonnet macaques are characterized not by immediate tit-for-tat reciprocal altruism, but by reciprocity over a longer time span, and that affiliative social relationships may be less constrained by agonistic relationships than is the case in more despotic species of macaques.     

Patterns of dominance and affiliation in wild pig-tailed macaques (Macaca nemestrina nemestrina) in West Sumatra

Patterns of agonistic and nonagonistic behaviors were studied in a troop of wild pig-tailed macaques in West Sumatra, Indonesia. The animals were provisioned and the identities of all adult and adolescent individuals were known. The females could be divided into high-, mid-, and low-ranking subsets of individuals. Most grooming occurred within, instead of between members of these subsets. The members of each subset also tended to feed together at the baiting sites, and they probably represented groups of close kin. Among females, grooming appeared to be a conciliatory behavior and was generally performed by the subordinate partner, while mounting was performed by the dominant partner and appeared to be a reassertion of dominance. High-ranking males tended to form agonistic alliances with females and to exchange grooming with estrous females. Low-ranking males did not have such associations with females and were frequently the targets of agonistic alliances of females and juveniles. Mounting between males appeared to be a conciliatory behavior. It seemed effective since severe aggression between males was not observed. The subordinate partner mounted more frequently than the dominant one, but the direction of mounting was apparently controlled by the latter. This suggests that, among pig-tailed macaques, the dominant male plays an important role in the coexistence of males in the troop.     

Serotonin and octopamine elicit stereotypical agonistic behaviors in the squat lobster Munida quadrispina (Anomura, Galatheidae)

Serotonin and octopamine have been implicated as modulators of posture and behavior in several crustaceans. Here we characterize the agonistic behaviors of normally interacting squat lobsters Munida quadrispina (Anomura, Galatheidae) and their responses to serotonin and octopamine injected into the ventral hemolymph sinus, in order to evaluate the potential roles of these amines in modulating agonistic behaviors. Normally interacting M. quadrispina do not develop lasting dominance hierarchies, although transient aggressive and submissive displays do occur. Injected serotonin elicits postures and behaviors in isolated individuals similar to those typical of aggressive, normally interacting animals. Injected octopamine can produce postures and behaviors typical of submissive animals, and elicits behaviors which imply a modulatory role for octopamine in tailflipping. The effects of both amines are reversible and dose dependent, and the dose-response curves parallel the normal progression of agonistic interactions. The social behaviors and reactions to injected serotonin and octopamine of M. quadrispina differ from those of lobsters and crayfish, indicating that interspecific differences in neuromodulation of behavior and motor output exist. Such differences have implications for the understanding of aminergic modulation of aggression and the evolution of aminergic modulation in crustaceans. Accepted: 22 June 1997     

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.     

Effect of testosterone and melatonin on social dominance and agonistic behavior in male Tscheskia triton

Social dominance and agonistic behavior play important roles in animal societies. Melatonin and testosterone are closely related to social dominance and agonistic behavior in rodents, but interactions between both of them remain unknown. In this study we investigated the effects of testosterone and melatonin by manipulating photoperiod and castration on social dominance and agonistic behavior in male Tscheskia triton. Castration significantly decreases social dominance of both short- and long-day males, suggesting that testosterone benefits social dominance of males in both breeding and non-breeding seasons. In intact conditions, long-day males tended to dominate short-day males, suggesting that the effect of testosterone on social dominance was a little stronger than melatonin. However, castrated short-day males became dominant over their castrated long-day opponents meaning that high melatonin levels obviously benefit social dominance in males. Hormone implantation indicated that testosterone had no effect on non-breeding condition, but that melatonin was important during the breeding season. Our results indicate that both testosterone and melatonin are important in determining social dominance in male hamsters, and the effect of testosterone appears to be stronger than melatonin. Testosterone is responsible for aggression and social dominance in male hamsters during the breeding season, while melatonin regulates behavior during non-breeding, probably due to the different seasonal secretory patterns of the hormones.     

Agonistic Behavior in the Squid Loligo plei (Loliginidae,Teuthoidea): Fighting Tactics and the Effects of Size and Resource Value

Observations of small schools of squids in captivity suggested that dominance relationships among males were based upon major differences in the frequency or duration of their agonistic behavior, but staged contests showed few differences. During staged contests, squids exhibited up to 21 separate behaviors. Some contests included a complex array of visual signals and side-by-side posturing (Lateral Display) followed by physical contact during Fin beating. There was behavioral variability and step-wise escalation during the contests: squids performed either 1. long sequences of visual signaling followed by Chasing and Fleeing; or 2. short sequences of visual signaling followed by physical Fin beating and ending with Chasing and Fleeing. Size influenced outcome in all contests; larger males were more likely to win the contest. Size had no effect on contest duration, but contest duration was shorter when resource value was high, especially when a male established temporary ownership of a female. We speculate that when the perceived resource value is high, male squids are more likely to engage in a shorter yet riskier fighting tactic.     

Direct benefits of social dominance in juvenile crayfish

Crayfish are known for their innate aggressiveness and willingness to quickly establish dominance relationships among group members. Consequently, the formation of dominance hierarchies and the analysis of behavioral patterns displayed during agonistic encounters have mostly been tested in environments that provide no immediate resources besides space. We tested the hypothesis that social hierarchy formation in crayfish serves to determine access to future resources. Individuals within groups of three juvenile crayfish were allowed to form a social hierarchy in a featureless environment before a single food resource was presented. Higher dominance indices were significantly correlated with increased access to the food. The highest ranked crayfish spent more time in contact with the food than did medium-ranked and lowest ranked crayfish, and crayfish of medium rank spent more time in contact with the resource than did lowest ranked animals. The highest ranked crayfish consolidated their dominant status in the presence of food, indicated by a complete absence of any submissive behaviors during that period. The results of these experiments show that the disposition of crayfish to engage in fighting and formation of a dominance hierarchy in a featureless environment serves to determine future access to an emerging resource, thereby entailing greater benefits for animals of higher social rank.     

Dominance rank reversals and rank instability among male Lemur catta: the effects of female behavior and ejaculation

In this study, dominance rank instability among male Lemur catta during mating was investigated. Also, data on agonism and sexual behavior across five consecutive mating seasons in a population of L. catta on St. Catherines Island, USA, were collected. Instances of male rank instability were categorized into three types. Type 1 consisted of a temporary switch in the dominance ranks of two males, which lasted for a period of minutes or hours. Type 2 dyadic male agonistic interactions showed highly variable outcomes for a period of time during which wins and losses were neither predictable nor consistent. Type 3 interactions consisted of a single agonistic win by a lower-ranked male over a more dominant male. More Type 2 interactions (indicating greater dominance instability) occurred when males had not spent the previous mating season in the same group, but this trend was not statistically significant. The majority of periods of male rank instability were preceded by female proceptivity or receptivity directed to a lower-ranked male. As such, exhibition of female mate choice for a lower-ranking male appeared to incite male-male competition. Following receipt of female proceptivity or receptivity, males who were lower-ranking took significantly longer to achieve their first agonistic win over a more dominant male than did males who were higher-ranked. Ejaculation frequently preceded loss of dominance. In conclusion, temporary rank reversals and overall dominance rank instability commonly occur among male L. catta in mating contexts, and these temporary increases in dominance status appear to positively affect male mating success.     

Developmental changes in the nature and function of social dominance among young children

Primate models of group social organization stress that dominant individuals receive more positive social behavior than other group members [Chance:Man 2:503–518 (1967); Seyfarth: Journal of Theoretical Biology 65:671–698 (1977)]. Such views suggest that short-term functions of dominance include the regulation of cohesive behavior, as well as the control of group conflict. However, this model has not been examined from a developmental perspective. In the present study, ten groups of children, who ranged in age from 8 to 66 months, were observed using both focal individual and matrix completion procedures. Social sequences were described according to a preestablished taxonomy of agonistic and affiliative actions. Agonistic exchanges ending with submission revealed linear dominance hierarchies at all age levels. In each group, certain children attracted more affiliative activities, and thus appeared as more popular individuals. However, affiliative gestures were distributed according to dominance status only at the two older age levels. Findings indicate that social dominance is developmentally the earliest stable dimension of peer group social organization and that cohesive activities are coordinated with dominance status toward the end of the preschool period.     

Differences in Behavior and Social Organization of Female Guinea Pigs as a Function of the Presence of a Male

Object of the study was the variability of social behavior and organization in female guinea pigs as a function of the presence of a male. Four groups of 8 females were observed. Two of these groups were without a male, the other two had a sexually experienced, sterilized male. All females were about 20 d old at the beginning of the study. In the groups without a male the females formed clear and stable dominance relationships resulting in a linear rank order. The frequency of agonistic behavior was higher compared to the other groups. Some of these females displayed male-like courtship behavior. Among the females in the groups with a male only the alpha and omega positions were stable, while the dominance relations between the other females remained unstable. These females directed about 40–50 % of their contact behavior towards the male. In relation to the presence or absence of a male the females differed in their social orientation: those who lived with a male were strongly orientated towards it, those living without a male were strongly oriented towards each other. The hypothesis is put forward, that the females in the two situations differed in their motivational states. For the females living with a male this animal was a stable point of social reference, while for the females living without a male their stable dominance structure — with more agonistic behavior — was the most important stabilizing factor.     

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.