On dominance relations and the structure of animal societies: I. Effect of inherent characteristics

Societies are considered in which a non-transitive dominance relation exists between every pair of members, such as the peck-right in a flock of hens. A one-dimensional measure of the structure of such a society,h, is defined, withh=0 for equality andh=1 for the hierarchy. It is assumed that each member of the society is characterized by an ability vector whose components depend on individual characteristics such as size, concentration of sex hormone, etc., but not on social factors such as social rank. The distribution of abilities among members of the society is assumed to be given by a distribution function which is the same for all members, and the probability that one member dominates another is given by a function of the ability vectors of the two. On these assumptions formulas for the expected (mean) value and variance ofh are determined in terms of the distribution and dominance probability functions. Some special cases are calculated, especially that for normany distributed abilities and dominance probability given by the normal probability integral. Several conclusions are derived. If all members are of equal ability, so that dominance probability is 1/2, then any sizable society is much more likely to be near the equality than the hierarchy; and, as the size of the society increases, the probability that it will be near the hierarchy becomes vanishingly small. If the dominance probability is a weighted sum of several independent components, which make up the ability vector, then the society is less likely to be close to the hierarchy as the number of these components increases. The hierarchy is the prevalent structure only if unreasonably small differences in ability are decisive for dominance. From this it appears that the social factors, or psychological factors such as the previous history of dominance, which are not included in the present treatment, may be of great importance in explaining the observed prevalence of structures very close to the hierarchy in flocks of domestic hens.     

On dominance relations and the structure of animal societies: II. Some effects of possible social factors

In a previous paper (Landau, 1951) it was shown that a society with a dominance relation would rarely tend to be close to the hierarchy in structure if dominance is determined solely by the inherent characteristics of the members. Here we consider the effects of other factors, due to social rank or to the outcome of previous encounters which affected dominance. The following results are obtained. A uniform bias against reversal of dominance will have no effect on the stationary distribution of the structure of the society. If the probability of dominance is a linear function of the previously established score (number of members dominated), there will be a small tendency for the society to move toward the hierarchy; but this is negligible for large societies. If a member never challenges another whose score exceeds his own by two or more, or if he can never dominate if he should challenge, then the hierarchy is the only stable structure. From the last result it is concluded that social factors which restrict challenges or the probability of dominance could easily account for societies close to the hierarchy, such as are observed in flocks of domestic hens. The effectiveness of social bias in establishing hierarchies is much greater in small societies than in large ones.     

Dominance hierarchy among workers changes with colony development in Polistes japonicus (Hymenoptera, Vespidae) paper wasp colonies with a small number of workers

Dominance hierarchy in the primitively eusocial wasp Polistes japonicus was analysed in four colonies for two periods: (1) the first-brood period, when only early emerging workers are present on the nest, and (2) the mixed-brood period, when the first and second (last) broods are present on the nest. The rank in the dominance hierarchy was determined based on a sociogram showing a dominance–subordinance relationship for all pairs of workers. During the first-brood period, older workers were likely to be more dominant (older dominance hierarchy), while the rank of workers was reversed during the mixed-brood period, with younger workers being likely to be more dominant (younger dominance hierarchy). However, the oldest and youngest workers were not always the top-ranked workers in the dominance hierarchy during the first- and mixed-brood periods, respectively, and during the mixed-brood period no younger dominance hierarchy was evident when the first or second brood was analysed separately. Higher ranked workers displayed dominance behaviour more frequently, and the lowest ranked worker hardly displayed dominance behaviour. Most workers displayed dominance behaviours primarily towards the worker ranked immediately below in the dominance hierarchy during the mixed-brood period but not during the first-brood period. The bodies of younger workers were larger for the mixed brood, but not for the first brood in some colonies or the second brood in all colonies. The association between body size and rank in the dominance hierarchy was negative during the first-brood period and positive during the mixed-brood period, with a nearly significant trend also seen even when the analysis was limited to the second brood. To explain the above temporal change from an older dominance hierarchy to a younger dominance hierarchy, we propose the hypothesis that the probability of a worker inheriting the colony increases rapidly with colony development, and consequently younger larger workers attempt to move up the dominance hierarchy in order to produce their own offspring by becoming the superseder late in colony development, rather than working harmoniously so as to boost the overall production of reproductive progeny for a colony, which is the strategy adopted early in colony development.     

On dominance relations and the structure of animal societies: III The condition for a score structure

The necessary and sufficient condition is given forn integers to be the score structure of a society with a dominance relation. A proof is also given for a theorem showing that there are members who dominate every other member either directly or indirectly through a single intermediate member.     

Monkeys in the Middle: Parasite Transmission through the Social Network of a Wild Primate

In wildlife populations, group-living is thought to increase the probability of parasite transmission because contact rates increase at high host densities. Physical contact, such as social grooming, is an important component of group structure, but it can also increase the risk of exposure to infection for individuals because it provides a mechanism for transmission of potentially pathogenic organisms. Living in groups can also create variation in susceptibility to infection among individuals because circulating levels of immunosuppressive hormones like glucocorticoids often depend on an individual’s position within the group’s social structure. Yet, little is known about the relative roles of socially mediated exposure versus susceptibility in parasite transmission among free-living animal groups. To address this issue, we investigate the relationship between host dominance hierarchy and nematode parasite transmission among females in a wild group of Japanese macaques (Macaca fuscata yakui). We use social network analysis to describe each individual female’s position within the grooming network in relation to dominance rank and relative levels of infection. Our results suggest that the number of directly-transmitted parasite species infecting each female, and the relative amount of transmission stages that one of these species sheds in faeces, both increase with dominance rank. Female centrality within the network, which shows positive associations with dominance hierarchy, is also positively associated with infection by certain parasite species, suggesting that the measured rank-bias in transmission may reflect variation in exposure rather than susceptibility. This is supported by the lack of a clear relationship between rank and faecal cortisol, as an indicator of stress, in a subset of these females. Thus, socially mediated exposure appears to be important for direct transmission of nematode parasites, lending support to the idea that a classical fitness trade-off inherent to living in groups can exist.     

Fight for your breeding right: hierarchy re-establishment predicts aggression in a social queue

Social aggression is one of the most conspicuous features of animal societies, yet little is known about the causes of individual variation in aggression within social hierarchies. Recent theory suggests that when individuals form queues for breeding, variation in social aggression by non-breeding group members is related to their probability of inheriting breeding status. However, levels of aggression could also vary as a temporary response to changes in the hierarchy, with individuals becoming more aggressive as they ascend in rank, in order to re-establish dominance relationships. Using the group-living fish, Neolamprologus pulcher, we show that subordinates became more aggressive after they ascended in rank. Female ascenders exhibited more rapid increases in aggression than males, and the increased aggression was primarily directed towards group members of adjacent rather than non-adjacent rank, suggesting that social aggression was related to conflict over rank. Elevated aggression by ascenders was not sustained over time, there was no relationship between rank and aggression in stable groups, and aggression given by ascenders was not sex-biased. Together, these results suggest that the need to re-establish dominance relationships following rank ascension is an important determinant of variation in aggression in animal societies.     

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.     

Hierarchical structure and the influence of individual attributes in the captive squirrel monkey (<Emphasis Type="Italic">Saimiri collinsi</Emphasis>)

The dominance structure of primate social groups varies widely. In addition to the groups’ composition, intrinsic attributes such as sex, body size and life experience are important factors that can affect hierarchical dominance relations. All primates are social animals, and the social environment has a direct influence on the physiological conditions of vital systems such as immunological, reproductive and cardiovascular systems. In this study, we analyze the hierarchical structure of Saimiri collinsi in captivity, including the hierarchical structure type, the influence of individual intrinsic characteristics (sex, age, weight and origin—born in captivity or in the wild) based on the prior-attributes model, the relation between agonistic behavior frequency and hierarchical position, and hierarchy steepness, which represents the dominance gradient. We found that the group order was characterized by a partial hierarchy: a dominance position could be occupied by more than one individual simultaneously, including individuals of both sexes. Intrinsic characteristics had no influence on hierarchical structure, with the exception of the male in the highest hierarchical position, which had a markedly larger body than all other group members. Thus, the prior-attributes model did not apply to hierarchical formation of S. collinsi in captivity. Only the frequency of agonistic behavior of males correlated with their hierarchical position, and they differed from all other group members in their more aggressive behavior. The steepness between adjacent positions along the dominance gradient was significant only between the dominant male and the next individual in the group, with a smooth gradient between the other positions in the rank. As the access to resources is directly related to hierarchical dominance, a smooth dominance gradient is to be expected in species that form very large groups, such as wild Saimiri populations.     

Sex differences in the effect of social status on the growth of subordinates in a co-operatively breeding cichlid

Social influences on the growth of the co-operatively breeding cichlid Neolamprologus pulcher were studied by examining the size structure of existing laboratory groups and the responses of males and females to removals of higher-ranked fish, which created larger size differences with more dominant fish. The size differences between adjacently ranked group members were predicted to differ from expected based on random size distributions of group members. Both males and females were predicted to respond to removals by increasing growth rate. In previously established groups, the size difference between dominant and highest-ranked subordinate males was greater than expected based on random group-assembly rules. The size difference between dominant and subordinate females did not differ from the null expectation. Third-ranked subordinate males increased their growth rate upon moving up one place in the dominance hierarchy (after the removal of higher-ranking fish) relative to fish that did not change rank. Contrary to predictions, however, females did not increase growth upon increasing rank.     

The Maintenance of Stable Dominance Hierarchies and the Pattern of Aggression: Support for the Suppression Hypothesis

Three hypotheses have been proposed to explain the development and maintenance of dominance hierarchies. According to the first hypothesis the dominance hierarchy is a result of the animals fighting once at their first encounter and then using the outcome of that fight to determine the rank order. The second hypothesis proposes that a dominance hierarchy reflects the fighting ability of the individuals in the group at each moment and is therefore relatively fluid with individuals continuously fighting for position. A third hypothesis, the suppression hypothesis, states that the dominance hierarchy is based to a large extent on the outcome of the first fight between the individuals but the dominant animal in each pair continuously attacks the subdominant individuals to condition them to lose in future encounters. We studied six well‐established flocks containing six adult hens each (Gallus gallus domesticus). Five of the flocks had linear hierarchies. The aggression was significantly more often directed towards the next low‐ranking individual. There was a good correlation between rank and comb size (height × width), but no significant correlation between rank and weight, or rank and level of fluctuating asymmetry. There was no significant correlation between levels of aggression and similarity of comb size for individuals of neighboring ranks. Our results tentatively support the suppression hypothesis for the maintenance of dominance hierarchies in the domestic hen.     

Dominance and queen succession in captive colonies of the eusocial naked mole-rat,Heterocephalus glaber.

Naked mole-rat colonies exhibit a high reproductive skew, breeding being typically restricted to one female (the ''queen'') and one to three males. Other colony members are reproductively suppressed, although this suppression can be reversed following the removal or death of the queen. We examined dominance and queen succession within captive colonies to investigate the relationship between urinary testosterone and cortisol, dominance rank and reproductive status; and to determine if behavioural and/or physiological parameters can be used as predictors of queen succession. Social structure was characterized by a linear dominance hierarchy before and after queen removal. Prior to queen removal, dominance rank was negatively correlated with body weight and urinary testosterone and cortisol titres in males and females. Queen removal results in social instability and aggression between high ranking individuals. Dominance rank appears to be a good predictor of reproductive status: queens are the highest ranking colony females and are succeeded by the next highest ranking females. The intense dominance-related aggression that accompanies reproductive succession in naked mole-rats provides empirical support for optimal skew theory.     

Dihydrotestosterone propionate effects on dominance and sexual behaviors in gonadectomized male and female rhesus monkeys.

One triad of male and two triads of female gonadectomized rhesus monkeys were observed as social groups assembled for repeated hour-long sessions. Social relationships were measured in terms of aggressive behavior between the members of each group in order to determine the dominance hierarchical order. Sexual performance was assessed for each male, before and after castration, in tests with an estrogen-stimulated ovariectomized female. Similar measures were made when the same female was periodically introduced to the all-male triad. When dihydrotestosterone propionate (DHTP) was administered for a period of 6 weeks to the middle-ranking member of each group, social status changes occurred in two groups, one male and one female, resulting in the elevation of the treated monkeys to the highest rank in the dominance hierarchy. In the other female group, aggressive behavior was increased with DHTP treatment of the middle-ranking female. Somatic effects, particularly a gain in body weight, occurred in all treated animals. Yawning behavior also increased significantly in those animals receiving DHTP. The latter two effects returned toward pretreatment levels following the cessation of hormone injection; however, changes in dominance hierarchy persisted to the end of the experiment, 6 weeks following the last DHTP treatment.     

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

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

Dominance hierarchies among communally held juvenile lobsters,Homarus americanus

An investigation was conducted to determine the time for establishment of dominance hierarchies, their stability, and their relationship to molting and death of cultured juvenile lobsters, Homarus americanus. Agonistic displays between communally held animals were monitored daily. Interactions between all possible pairs of animals within each group were used to determine formation of a dominance hierarchy. A decrease in the frequency of agonistic interactions was recorded after the first day of the experiment. A stable four‐rank hierarchy was established within 2–3 days. When the dominant animal was molting, the subordinates displayed aggressively toward all members of the group but remained submissive to the dominant. When the dominant animal died, stability was lost and aggressive interactions among subordinates established a new ranking. Molting subordinates were highly susceptible to injury and death. Loss of subordinates did not affect the ranking of individuals.     

The changes in the dominance hierarchy over time of a complete field-captured colony of Cryptomys hottentotus hottentotus

The common mole-rat, Cryptomys h. hottentotus , is a social subterranean rodent occurring in colonies in which one female and one to three males are involved in reproduction and the remaining colony members are non-reproductive. Within each sex the reproductive animals are usually the largest and most dominant animals.
The dominance hierarchy amongst a field-captured colony was linear ( h = 0.95, calculated from Landau's linearity index) soon after capture. The non-reproductive females were ranked low in the dominance hierarchy; many were subordinate to non-reproductive males. The order of capture of mole-rats was not related to the position in the dominance hierarchy. The hierarchy became non-linear ( h = 0.56) after six months in captivity during which two juvenile animals became adult. The breakdown in the hierarchy may result from the lack of opportunity in captivity for animals to disperse and establish satellite colonies, or from colony members becoming co-dominant in the hierarchy as a result of a rise in rank by young animals.
Dominant mole-rats are involved in a greater proportion of interactive behaviours than subordinates. Popularity studies show that females tend to be more popular animals than males. The largest reproductive male was the least popular animal in the first study, whereas a beta male was the least popular animal in the second study period. The reproductive female was the most popular in both periods.     

Genealogical and cross-genealogical dominance relations in a group of free-ranging rhesus monkeys (Macaca mulatta) on Cayo Santiago

Observations of dominance relations in a large group of rhesus monkeys on Cayo Santiago were carried out over a period of 25 months. Dyadic interactions in which an aggressive gesture in one individual was followed by a submissive gesture in the other were recorded as fights and considered reliable indices of dominance. The analysis revealed the following characteristics: (1) Maternal dominance over female offspring; (2) maternal dominance over male offspring up the age six years at which time the son leaves his natal group or remains in the group and rises in rank over his mother; (3) dominance of older brothers over younger male siblings until the age of five years at which time the younger brother rises in rank; (4) rank reversal between sisters when the younger sister reaches the age of three to four years; (5) brother-sister relative rank dependency on age until the male sibling reaches three to four years at which time he rises in rank; (6) linear dominance relations in the crossgenealogical dominance hierarchy; and (7) linear, but unstable, dominance relations in the adult male hierarchy. With few exceptions, the pattern of genealogical, cross-genealogical, and adult male dominance relations in the group under study was consistent with data reported for a small social group (group F) on Cayo Santiago and for Japanese macaques.     

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

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

Outline of a probabilistic approach to animal sociology: III

The probabilities of the emergence of the two kinds of social structure in a 3-bird flock (chain and cycle) are deduced under the assumption of certain biases acting on the social dynamics of the flock. In particular a bias against the reversal of peck order and a bias against encounters of individuals of disparate social rank are considered. Like-wise a distribution of an “inherent” fighting ability is considered which influences the outcomes of encounters. A functional relation is derived between the importance of this ability and the initial probability of a chain structure.     

Interrelationship of serum testosterone, dominance and ovarian cyclicity status in female African elephants

The African elephant population in North American zoos is not self-sustaining, in part due to the prevalence of ovarian acyclicity. While little is known about the cause of this condition, earlier research has shown that females without cyclic corpus luteum (CL) function rank higher in the dominance hierarchy than females with cyclic CL function. The goal of this study was to measure longitudinal serum testosterone concentrations in captive female African elephants to determine if there is a relationship among serum testosterone concentrations, social dominance rank and ovarian cyclicity status. Weekly blood samples from 49 female African elephants (24 having and 25 not having cyclic CL function at 22 facilities) were collected over a 12-month period and analyzed for serum testosterone using an enzymeimmunoassay. A progesterone radioimmunoassay was used to quantify serum progestagen concentrations and categorize ovarian cyclicity status. The dominance hierarchy of individual elephants within each herd was assessed by a written temperament survey, which identified 19 dominant, 15 middle and 15 subordinate females. No clear patterns of serum testosterone secretion were observed in females with and without cyclic CL function. Furthermore, no significant relationships were found among serum testosterone concentrations, dominance rank, and ovarian cyclicity status. These data suggest that increased circulating testosterone concentrations are not associated with greater rates of ovarian acyclicity or dominance status in captive female African elephants.     

Knowledge acquired and decisions made: triadic interactions during allogrooming in wild bonnet macaques,Macaca radiata.

The pressures of developing and maintaining intricate social relationships may have led to the evolution of enhanced cognitive abilities in many nonhuman primates. Knowledge of the dominance ranks and social relationships of other individuals, in particular, is important in evaluating one''s position in the rank hierarchy and affiliative networks. Triadic interactions offer an excellent opportunity to examine whether decisions are taken by individuals on the basis of such knowledge. Allogrooming supplants among wild female bonnet macaques (macaca radiata) usually involved the subordinate female of a grooming dyad retreating at the approach of a female dominant to both members of the dyad. In a few exceptional cases, however, the dominant member of the dyad retreated; simple non-cognitive hypotheses involving dyadic rank differences and agonistic relationships failed to explain this phenomenon. Instead, retreat by the dominant individual was positively correlated with the social attractiveness of her subordinate companion (as measured by the duration of grooming received by the latter from other females in the troop). This suggests that not only does an individual evaluate relationships among other females, but does so on the basis of the amount of grooming received by them. Similarly, the frequency of approaches received by any female was correlated with her social attractiveness when she was the dominant member of the dyad, but not when she was the subordinate. This indicated that approaching females might be aware of the relative dominance ranks of the two allogrooming individuals. In logistic regression analyses, the probability of any individual retreating was found to be influenced more by her knowledge of her rank difference with both the other interactants, rather than by their absolute ranks. Moreover, information about social attractiveness appeared to be used in terms of correlated dominance ranks. The nature of knowledge acquired by bonnet macaque females may thus be egotistical in that other individuals are evaluated relative to oneself, integrative in that information about all other interactants is used simultaneously, and hierarchical in the ability to preferentially use certain categories of knowledge for the storage of related information from other domains.