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.     

A Novel Method of Assessing Dominance Hierarchies Shows Nuance,Linearity and Stability in the Dinosaur Ant Dinoponera quadriceps

Many social species with relatively simple societies have dominance hierarchies of individuals, with dominant individuals achieving fitness and subordinate individuals either queuing to obtain fitness or achieving only indirect fitness by helping relatives. Assessing the dominance hierarchy in a social group is generally based upon observing dyadic interactions as and when they occur spontaneously within the whole‐group setting. However, this method can be very time‐consuming because many dyads interact only very rarely, necessitating either extremely long observation periods or many dyadic relationships being unresolved. Here, we report an alternative method using the queenless dinosaur ant Dinoponera quadriceps, which lives in colonies containing tens of individuals. We removed all individuals from their nest and observed the dominance behaviours expressed in isolated dyadic interactions for every pairwise combination of individuals. Individuals showed a classic dominance behaviour in this setting, and the rapid nature of the assay allowed us to observe every dyadic relationship on a weekly basis over 4 weeks. The dominance hierarchies based on these isolated dyadic interactions correlated well with those produced by the conventional method of colony observations. They showed the hierarchies to be highly linear and stable, and also revealed that dominance relationships may extend further down the hierarchy than previously thought. Although highly manipulative, the isolated dyadic interaction method works well and will likely make more feasible the study of other social species in which pairs of individuals can be isolated together.     

Effect of unknown relationships on linearity, steepness and rank ordering of dominance hierarchies: simulation studies based on data from wild monkeys

The presence of unknown dyadic relationships is a common problem in constructing dominance hierarchies for groups of social animals. Although previously acknowledged, the influence of unknown relationships on hierarchy measures like linearity and steepness has not been studied in detail. Using real data-sets from four groups of wild monkeys, we illustrate how unknown relationships affect linearity and steepness of hierarchies and the consistency of rank ordering based on de Vries’ I&SI method. Monte Carlo simulations revealed significant negative linear relationships between the proportion of unknown relationships and both linearity and steepness. These simulations over-estimated steepness and linearity indices relative to additional real-data input matrices. Rank orders became inconsistent at 26–38% unknown relationships, depending on the group. Group size and the specific input matrix substantially affected how much unknown relationships influenced steepness and linearity, the values of these indices and the point at which rank order became inconsistent. We recommend caution in characterizing the dominance structure of a group with many unknown relationships, and in drawing conclusions about hierarchy linearity and steepness based on few input matrices, especially if they contain many unknown relationships. Quantitative characterizations of hierarchies are perhaps best viewed as a somewhat fluid range rather than fixed values.     

The ghost of social environments past: dominance relationships include current interactions and experience carried over from previous groups

Dominance hierarchies pervade animal societies. Within a static social environment, in which group size and composition are unchanged, an individual's hierarchy rank results from intrinsic (e.g. body size) and extrinsic (e.g. previous experiences) factors. Little is known, however, about how dominance relationships are formed and maintained when group size and composition are dynamic. Using a fusion-fission protocol, we fused groups of previously isolated shore crabs (Carcinus maenas) into larger groups, and then restored groups to their original size and composition. Pre-fusion hierarchies formed independently of individuals' sizes, and were maintained within a static group via winner/loser effects. Post-fusion hierarchies differed from pre-fusion ones; losing fights during fusion led to a decline in an individual's rank between pre- and post-fusion conditions, while spending time being aggressive during fusion led to an improvement in rank. In post-fusion tanks, larger individuals achieved better ranks than smaller individuals. In conclusion, dominance hierarchies in crabs represent a complex combination of intrinsic and extrinsic factors, in which experiences from previous groups can carry over to affect current competitive interactions.     

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

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

On the Additive and Dominant Variance and Covariance of Individuals Within the Genomic Selection Scope

Genomic evaluation models can fit additive and dominant SNP effects. Under quantitative genetics theory, additive or “breeding” values of individuals are generated by substitution effects, which involve both “biological” additive and dominant effects of the markers. Dominance deviations include only a portion of the biological dominant effects of the markers. Additive variance includes variation due to the additive and dominant effects of the markers. We describe a matrix of dominant genomic relationships across individuals, D, which is similar to the G matrix used in genomic best linear unbiased prediction. This matrix can be used in a mixed-model context for genomic evaluations or to estimate dominant and additive variances in the population. From the “genotypic” value of individuals, an alternative parameterization defines additive and dominance as the parts attributable to the additive and dominant effect of the markers. This approach underestimates the additive genetic variance and overestimates the dominance variance. Transforming the variances from one model into the other is trivial if the distribution of allelic frequencies is known. We illustrate these results with mouse data (four traits, 1884 mice, and 10,946 markers) and simulated data (2100 individuals and 10,000 markers). Variance components were estimated correctly in the model, considering breeding values and dominance deviations. For the model considering genotypic values, the inclusion of dominant effects biased the estimate of additive variance. Genomic models were more accurate for the estimation of variance components than their pedigree-based counterparts.     

Dominance hierarchies in male lizards: Implications for zoo management programs

Depending on spatial requirements and the distribution of key resources in the environment, social behavior among lizards varies from defense of exclusive territories to the establishment of dominance hierarchies. In captivity or under conditions where dispersal is not possible, dominance hierarchies often emerge in species that are otherwise territorial. This review explores some of the morphological, behavioral, and hormonal determinants of social status in male lizards and how these may lead to differential reproductive function in dominant and subordinate individuals. Emphasis is placed on the importance of population density, local resource dispersion, and the composition and stability of social groups in promoting hierarchical behavior. Results of these studies have ramifications for several aspects of zoo management, including exhibit design, choice of animals to be housed together, provision of resources in space and time, and orientation of enclosures within captive breeding facilities. © 1994 Wiley-Liss, Inc.     

A unified model of dominance hierarchy formation and maintenance

In many different species it is common for animals to spend large portions of their lives in groups. Such groups need to divide available resources amongst the individuals they contain and this is often achieved by means of a dominance hierarchy. Sometimes hierarchies are stable over a long period of time and new individuals slot into pre-determined positions, but there are many situations where this is not so and a hierarchy is formed out of a group of individuals meeting for the first time. There are several different models both of the formation of such dominance hierarchies and of already existing hierarchies. These models often treat the two phases as entirely separate, whereas in reality, if there is a genuine formation phase to the hierarchy, behaviour in this phase will be governed by the rewards available, which in turn depends upon how the hierarchy operates once it has been formed. This paper describes a method of unifying models of these two distinct phases, assuming that the hierarchy formed is stable. In particular a framework is introduced which allows a variety of different models of each of the two parts to be used in conjunction with each other, thus enabling a wide range of situations to be modelled. Some examples are given to show how this works in practice.     

非人灵长类雌性等级的研究方法

在非人灵长类动物等级研究中, 有关判定与描述优势等级方法的选择至关重要。早期对旧大陆灵长类的等级研究主要关注雄性, 对雌性涉猎较少。目前主要应用攻击-屈服、取食、携婴、相互理毛等行为标准来研究灵长类的雌性等级, 也发现不同的物种往往有不同的雌性等级模式。依据社群结构的完整性将其分为线性等级与非线性等级, 并采用不同的方式进行描述。本文通过查阅文献资料, 对雌性等级的研究内容、判定标准以及描述方式进行论述, 以便抛砖引玉提高非人灵长类雌性等级的研究水平。     

A theory of leadership in human cooperative groups

Two types of models aim to account the origins of rank differentiation and social hierarchy in human societies. Conflict models suggest that the formation of social hierarchies is synonymous with the establishment of relationships of coercive social dominance and exploitation. Voluntary or 'integrative' models, on the other hand, suggest that rank differentiation - the differentiation of leader from follower, ruler from ruled, or state from subject - may sometimes be preferred over more egalitarian social arrangements as a solution to the challenges of life in social groups, such as conflict over resources, coordination failures, and free-riding in cooperative relationships. Little formal theoretical work, however, has established whether and under what conditions individuals would indeed prefer the establishment of more hierarchical relationships over more egalitarian alternatives. This paper provides an evolutionary game theoretical model for the acceptance of leadership in cooperative groups. We propose that the effort of a leader can reduce the likelihood that cooperation fails due to free-riding or coordination errors, and that under some circumstances, individuals would prefer to cooperate in a group under the supervision of a leader who receives a share of the group's productivity than to work in an unsupervised group. We suggest, in particular, that this becomes an optimal solution for individual decision makers when the number of group members required for collective action exceeds the maximum group size at which leaderless cooperation is viable.     

Female philopatry and dominance patterns in wild geladas

Cercopithecines have a highly conserved social structure with strong female bonds and stable, maternally inherited linear dominance hierarchies. This system has been ascribed to the pervasiveness of female philopatry within the typical multi-male, multi-female social groups. We examined the relationship between female philopatry, dominance hierarchies, and reproduction in geladas (Theropithecus gelada), a species with an unusual multi-leveled society. During a 4-year field study on a wild population in the Simien Mountains National Park, Ethiopia, we observed 14 units across two bands of geladas that underwent a number of events, such as male takeovers and female deaths, which could potentially disrupt female relationships and unit structure. First, we corroborate earlier reports that gelada females are natally philopatric: we observed no interunit migrations, and the female mortality rate was comparable to that of philopatric baboons (suggesting all female disappearances were indeed deaths). Second, contrary to previous reports, data from this long-term study show that geladas exhibit the linear and stable dominance hierarchies typical of other Cercopithecines. Moreover, female ranks appear to be maternally inherited. Third, we found no evidence that alpha females aggressively target the lowest ranking individuals, nor did rank confer clear reproductive advantages to dominant females within our 4-year observation period. As such, geladas fit the allostatic load model [Goymann & Wingfield, Animal Behaviour 67:591-602, 2004]. Our study confirms the importance of female philopatry in the kin-based Cercopithecine dominance system.     

Inferring dominance interactions from automatically recorded temporal data

Understanding dominance hierarchies is an important aspect of many studies of animal groups. Collecting the necessary data requires a large amount of time and effort to detect only a small number of interactions. Here, we assess the viability of extracting interactions from a temporal data stream of arrival and departures to a feeder equipped with a radio‐frequency identifier antenna. We show that the interactions extracted from this type of data can create similar dominance hierarchies to those produced by analysing video data of interactions. Additionally, we assessed the amount of temporal data required to obtain a dominance hierarchy and discuss the advantages and disadvantages of utilising temporal data streams to generate dominance hierarchies of animal groups. The code to extract the dominance interactions and test the correlation between different dominance hierarchies is included in the Supporting Information .     

Measuring Dominance and Constructing Hierarchies: An Example Using Mule Deer

Important issues that are still unresolved in the study of animal social groups are how dominance is measured and how individuals are ranked. Based on observations of mule deer (Odocoileus hemionus hemionus), we constructed hierarchies for the same 26 bucks using five potential correlates of dominance and three measures based on the outcome of agonistic interactions. Data for each of six behavior measures were converted to ranks by six different numerical techniques. Different measures of behavior resulted in different hierarchies. This was especially so when the hierarchy based on sparring rates was compared to hierarchies based on other criteria. Although our results showed that dominance is not a unitary concept, several measures of dominance were highly correlated with one another. Thus, one measure, in some instances, may be a good but imperfect predictor of dominance defined by other criteria. For data sets based on index scores or rates of performing behavior, the mean difference and standard-error difference ranking methods had distinct advantages. Both produced equally weighted ranks that minimized the effects of sampling errors.     

Effects of host social hierarchy on disease persistence

The effects of social hierarchy on population dynamics and epidemiology are examined through a model which contains a number of fundamental features of hierarchical systems, but is simple enough to allow analytical insight. In order to allow for differences in birth rates, contact rates and movement rates among different sets of individuals the population is first divided into subgroups representing levels in the hierarchy. Movement, representing dominance challenges, is allowed between any two levels, giving a completely connected network. The model includes hierarchical effects by introducing a set of dominance parameters which affect birth rates in each social level and movement rates between social levels, dependent upon their rank. Although natural hierarchies vary greatly in form, the skewing of contact patterns, introduced here through non-uniform dominance parameters, has marked effects on the spread of disease. A simple homogeneous mixing differential equation model of a disease with SI dynamics in a population subject to simple birth and death process is presented and it is shown that the hierarchical model tends to this as certain parameter regions are approached. Outside of these parameter regions correlations within the system give rise to deviations from the simple theory. A Gaussian moment closure scheme is developed which extends the homogeneous model in order to take account of correlations arising from the hierarchical structure, and it is shown that the results are in reasonable agreement with simulations across a range of parameters. This approach helps to elucidate the origin of hierarchical effects and shows that it may be straightforward to relate the correlations in the model to measurable quantities which could be used to determine the importance of hierarchical corrections. Overall, hierarchical effects decrease the levels of disease present in a given population compared to a homogeneous unstructured model, but show higher levels of disease than structured models with no hierarchy. The separation between these three models is greatest when the rate of dominance challenges is low, reducing mixing, and when the disease prevalence is low. This suggests that these effects will often need to be considered in models being used to examine the impact of control strategies where the low disease prevalence behaviour of a model is critical.     

Social Dominance Hierarchies and Phenotypic Correlates of Dominance in Captive Groups of the Grey‐cheeked Fulvetta Alcippe morrisonia in Taiwan

Dominance hierarchies usually form quickly among avian foraging groups because they are beneficial to most individuals by reducing conflict. Several characteristics that correlate with dominance rank have been identified in birds, but most of these conclusions rely on studies of temperate species. Hence, we studied whether captive group members of a subtropical species, grey‐cheeked fulvetta Alcippe morrisonia, form social dominance hierarchies when competing for food during the non‐breeding season. We also investigated whether sex, age, body condition and fat score were related to an individual's dominance rank which was established by counting aggressive interactions in six captive groups of nine individuals each. In all groups, linear dominance hierarchies were formed whereby yearlings dominated over adult birds, and individuals with a better body condition were also more dominant, while sex and fat score had no discernable effect. Male yearlings had significantly higher body masses and body condition indices than male adults, while female yearlings had significantly higher body masses, body condition indices and fat scores than female adults. However, there were no significant differences between male and female yearlings or adults for any of these variables. We suggest possible reasons for the dominance of yearlings, such as captive conditions or the higher body weight of yearlings.     

Allogrooming, partner choice, and dominance in male anubis baboons

This study was designed to test the hypothesis that among unrelated male baboons (Papio cynocephalus anubis) in single-gender social groups there is no significant association between dominance status and allogrooming performance. The hypothesis was tested using behavioral measures obtained by focal animal sampling techniques. The results indicate that unrelated male baboons established well-defined linear dominance hierarchies, formed allogrooming relationships with one another, and exhibited a nonrandom distribution of allogrooming; however, there were no significant relationships between dominance rank and the frequency of allogrooming. We further tested our results by grouping individuals into three dominance status classes (high, middle, and low) and comparing the classes. Analysis of variance demonstrated no significant differences in rates of allogrooming by dominance class. These results suggest that dominance did not account for the variation in observed allogrooming behavior: Dominance status did not appear to determine the frequency with which animals groomed others, the number of grooming partners, or frequency of grooming that any individual received in comparison to that performed. High-ranking animals did not have significantly more grooming partners than low-ranking animals, and there appeared to be little competition within the groups for subordinates to groom high-ranking animals. When age, kinship, and group tenure are controlled, performance and reception of allogrooming are not strongly associated with dominance in single-gender social groups of male anubis baboons.     

Dominance and association among members of a captive and a free-ranging group of grey kangaroos (Macropus giganteus)

Dominance relationships and the degree of association between members of groups of eastern grey kangaroos (Macropus giganteus) were investigated by observing a small group in captivity and a larger free-ranging group. Only decisive aggressive interactions were used as a criterion of dominance, and the number of times individuals occurred together were used as a measure of association. A dominance hierarchy in which the male assumed the highest rank was found in the captive group, while in the wild hierarchies were found separately among both males and females in the group. A low level of positive association existed between members of both the groups studied and the data indicated that in general the free ranging animals tended to move about at random with respect to each other. In the captive group dispersion about the enclosure was regular but became more random during the day in winter and spring.     

Can Individual Differences in History of Dominance Explain the Development of Linear Dominance Hierarchies?

Observed dominance hierarchies are often more linear than expected from randomly-formed dominance relationships, and in triads of animals attacks are distributed non-randomly. I hypothesize that an individual's history of dominance affects its probability of initiating aggressive interactions in the future and that individuals with winning records are more likely to initiate (winning begets initiating). Consistent with this hypothesis, evidence is presented that dominant individuals are more likely to attack than subordinate individuals. The winning begets initiating hypothesis may also explain why correlations between predicted dominance ranks (based on size, age etc.) and observed dominance ranks can be low: If the cost of engaging in and losing an interaction is high relative to the potential benefits of winning, then a large individual conditioned to be subordinate may refrain from contesting smaller, dominant individuals despite its actual competitive superiority.     

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.