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.     

Preservation of winter social dominance status in Brent Geese Branta bernicla bernicla within and across winters

Dominant and subordinate individuals in a group may benefit from the stability of the social dominance organisation, avoiding excessive waste of time and energy in aggressive interactions and reducing injury risks. Nevertheless, the likely evolutionary incentive for individuals to become, and furthermore to stay, dominant may destabilise such dominance hierarchies. In this context, the relative importance of fixed (e.g. sex, morphological size) and fluctuating (e.g. body condition, mating status, reproductive success, social unit size) traits influencing the establishment and preservation of dominance relationships could play a key role in group structure. We investigated the relative role of fixed and fluctuating traits on social status in Dark-bellied Brent Geese Branta bernicla bernicla which form large fairly unstable groups both within and across winters. We compared individual dominance scores of ringed Brent Geese during four consecutive winters. Brent Geese conserved their dominance score within a given winter irrespective of their age but were generally unable to conserve it across consecutive winters. As winter dominance scores correlated best with social unit size, dominance status thus appeared to be mostly a by-product of a fluctuating trait: breeding success in the previous summer. When we considered only adults that had the same social unit size during two consecutive winters, we observed a significant preservation of dominance scores. This result suggests that a fixed trait such as sex or morphological size may still play a role in setting dominance status.     

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.     

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.     

Choosing and using diversity indices: insights for ecological applications from the German Biodiversity Exploratories

Biodiversity, a multidimensional property of natural systems, is difficult to quantify partly because of the multitude of indices proposed for this purpose. Indices aim to describe general properties of communities that allow us to compare different regions, taxa, and trophic levels. Therefore, they are of fundamental importance for environmental monitoring and conservation, although there is no consensus about which indices are more appropriate and informative. We tested several common diversity indices in a range of simple to complex statistical analyses in order to determine whether some were better suited for certain analyses than others. We used data collected around the focal plant Plantago lanceolata on 60 temperate grassland plots embedded in an agricultural landscape to explore relationships between the common diversity indices of species richness (S), Shannon's diversity (H'), Simpson's diversity (D₁), Simpson's dominance (D₂), Simpson's evenness (E), and Berger–Parker dominance (BP). We calculated each of these indices for herbaceous plants, arbuscular mycorrhizal fungi, aboveground arthropods, belowground insect larvae, and P. lanceolata molecular and chemical diversity. Including these trait‐based measures of diversity allowed us to test whether or not they behaved similarly to the better studied species diversity. We used path analysis to determine whether compound indices detected more relationships between diversities of different organisms and traits than more basic indices. In the path models, more paths were significant when using H', even though all models except that with E were equally reliable. This demonstrates that while common diversity indices may appear interchangeable in simple analyses, when considering complex interactions, the choice of index can profoundly alter the interpretation of results. Data mining in order to identify the index producing the most significant results should be avoided, but simultaneously considering analyses using multiple indices can provide greater insight into the interactions in a system.     

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.     

Short-term exposure to testosterone propionate leads to rapid bill color and dominance changes in zebra finches

Testosterone (T) can influence both male-male competition and mate choice displays. In zebra finches, female mate choice is based in part on bill color, and bill color has been shown to be enhanced by long-term testosterone supplementation. However, it is not clear whether bill color plays a role in male-male interactions and how bill color responds to shorter-term changes in T. We tested whether a single injection of testosterone propionate (TP) would influence male-male dominance interactions and lead to rapid (over a three-day period) changes in bill color. In addition, we tested whether bill color predicted aggression and dominance. We allowed birds in triads to establish hierarchies and then injected either dominant or subordinate individuals with TP, in addition to establishing sham control triads. We found that red chroma, but not hue, predicted aggressiveness of males. Exposure to TP led both dominant and subordinate birds to increase dominance scores over three days, longer than the < 24 h period in which injected TP stays active. In addition, exposure to TP increased red chroma and hue in three days showing the dynamic nature of allocation of pigments to the bill. Our results suggest that zebra finches can modulate T and bill color levels over short time periods and these changes may occur through positive feedback between T-levels and dominance.     

Socio-spatial levels in linearity analysis of dominance hierarchies: a case study on elephant seals

The analysis of linearity is a key aspect of the study of dominance hierarchies. To study the effect of the choice of socio-spatial level of analysis, we calculated linearity in a large set of southern elephant seal (Mirounga leonina) hierarchies from two populations (Valdés Peninsula and Falkland Islands). The socio-spatial level of analysis affects the observational effort, the completeness of matrices, and the frequency of unknown relationships. These factors, in turn, have a notable effect on linearity. We conclude that dominance should be studied at the local level, where the absence of structural zeros and the low incidence of observational zeros produce complete matrices, well rooted in the true spatial and social structure of the population. Depending on the specific social system, the extrapolation of dominance from the local level to higher levels may result in sparse matrices, and in biased estimates of linearity. The variation of the socio-spatial level of analysis may in part explain the contrasting results obtained in different studies of linearity of dominance hierarchies. Electronic Publication     

Effects of anger on dominance-seeking and aggressive behaviors

Anger may have evolved to orchestrate social bargaining behaviors, which ultimately can lead to establishment of dominance hierarchies. Although the relationship between anger and dominance has strong empirical support, most studies have focused on visual cues of dominance. Across two experiments, we tested the hypothesis that anger increases dominance-seeking and agonistic behaviors in those who feel it. In the first experiment (n?=?82), we induced anger through a hostile mock debate and measured corrugator electromyographic activity, testosterone and cortisol levels, status-seeking tendency, and aggression using behavioral tasks. Compared with the control group, the anger group showed higher levels of aggression and status seeking, with the moderator effect of anger intensity. In the second experiment (n?=?162), anger, fear, sadness, and neutral state were induced by film clips, after which dominance-related behavioral tendencies were assessed. The anger group showed higher dominance scores, differing significantly from the fear, sadness, and/or control groups. These findings reinforce the notion that feelings of anger can cause an increase in status-seeking and agonistic behaviors, leading to possible action tendencies for the establishment of dominance hierarchies.     

Personality Predicts Social Dominance in Male Domestic Fowl

Individuals in social species commonly form dominance relationships, where dominant individuals enjoy greater access to resources compared to subordinates. A range of factors such as sex, age, body size and prior experiences has to varying degrees been observed to affect the social status an individual obtains. Recent work on animal personality (i.e. consistent variation in behavioural responses of individuals) demonstrates that personality can co-vary with social status, suggesting that also behavioural variation can play an important role in establishment of status. We investigated whether personality could predict the outcome of duels between pairs of morphologically matched male domestic fowl (Gallus gallus domesticus), a species where individuals readily form social hierarchies. We found that males that more quickly explored a novel arena, or remained vigilant for a longer period following the playback of a warning call were more likely to obtain a dominant position. These traits were uncorrelated to each other and were also uncorrelated to aggression during the initial part of the dominance-determining duel. Our results indicate that several behavioural traits independently play a role in the establishment of social status, which in turn can have implications for the reproductive success of different personality types.     

Social mediation of sexually selected ornamentation and steroid hormone levels in male junglefowl

Honest signals in sexual selection may be maintained by a variety of mechanisms. Comb size in male red junglefowl, Gallus gallus, a well-known predictor of female mate choice, is mediated by health and condition. Social status has also been shown to mediate comb size. To determine whether hormones related to male dominance behaviour might be related to social status and comb size, we monitored changes in plasma levels of testosterone and corticosterone following manipulation of social status. We removed young adult male junglefowl from their all-male flocks and placed them either in individual cages or in smaller flocks. We measured comb size and other morphological variables, as well as testosterone and corticosterone levels, before and after the manipulation of social status. Males in small flocks showed reduced comb growth, increased corticosterone and decreased testosterone levels compared with individually housed males. Within flocks, comb size was positively related to dominance rank and to testosterone level, although not always significantly. However, dominance rank was unrelated to either corticosterone or testosterone levels. In isolated birds, testosterone level was not related to comb size. Corticosterone level was not related to comb size in either treatment group. Our results are consistent with social mediation of comb size and testosterone and corticosterone levels, but it is unclear whether these hormones contribute to the observed social limitation of comb length.Copyright 2002 The Association for the Study of Animal Behaviour. Published by Elsevier Science Ltd. All rights reserved .     

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.     

Does testosterone determine dominance in the house sparrow Passer domesticus? An experimental test

Badges of status function in many birds within a social context to establish dominance hierarchies and reduce antagonistic encounters. In order to maintain the honesty of the signalling system, such badges must be costly to produce or to maintain. The chest bib of the house sparrow functions as a badge of status and changes in size are known to be controlled by testosterone levels. We sought to test the relative importance of testosterone as opposed to bib size in determining dominance within a group of male house sparrows. We did this by manipulating testosterone levels independently during both breeding and post‐breeding seasons in experimental birds and examining the effect of testosterone titre, as well as corticosterone titre relative to bib size on dominance levels. Dominance hierarchies within the groups were tested during both the breeding and post‐breeding phases. We compared the results of these tests with dominance among intact (unmanipulated) birds. Results suggested that the breeding season dominance levels were largely determined by testosterone levels as well as bib size, whereas the post‐breeding dominance levels were determined by post‐breeding testosterone titre and previous breeding season dominance level. Within unmanipulated birds, basal corticosterone levels were significantly, negatively correlated with dominance level, but only during the breeding season. The influence of breeding season dominance on post‐breeding dominance suggests social history is important in determining dominance interactions as well as current testosterone levels and bib size.     

Social context predicts recognition systems in ant queens

Recognition of group‐members is a key feature of sociality. Ants use chemical communication to discriminate nestmates from intruders, enhancing kin cooperation and preventing parasitism. The recognition code is embedded in their cuticular chemical profile, which typically varies between colonies. We predicted that ants might be capable of accurate recognition in unusual situations when few individuals interact repeatedly, as new colonies started by two to three queens. Individual recognition would be favoured by selection when queens establish dominance hierarchies, because repeated fights for dominance are costly; but it would not evolve in absence of hierarchies. We previously showed that Pachycondyla co‐founding queens, which form dominance hierarchies, have accurate individual recognition based on chemical cues. Here, we used the ant Lasius niger to test the null hypothesis that individual recognition does not occur when co‐founding queens do not establish dominance hierarchies. Indeed, L. niger queens show a similar level of aggression towards both co‐foundresses and intruders, indicating that they are unable of individual recognition, contrary to Pachycondyla. Additionally, the variation in chemical profiles of Lasius and Pachycondyla queens is comparable, thus informational constraints are unlikely to apply. We conclude that selection pressure from the social context is of crucial significance for the sophistication of recognition systems.     

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.     

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 reliability of palatability estimates obtained from rumen contents analysis and a field-based index of diet selection

Understanding herbivore foraging behaviour is often limited by not knowing the palatability of food items. Electivity indices can overcome this challenge by relating the consumption of food items to their relative abundance in the environment. We compare eight widely used electivity indices to relate the level of consumption of 48 food items in the rumen contents of 402 red deer Cervus elaphus in New Zealand to the environmental availability of food items. Additionally, we derive a field-based index of palatability from observed browse damage recorded during vegetation surveys at our site. Electivity indices were positively correlated with each other and with the browse survey index, which was also positively correlated with electivity indices from other studies. Thus, the choice of electivity index is unimportant when making comparisons of relative palatability among food items. Our analyses suggest that quantitative measures of palatability may be reliably inferred from observed browse damage in the field, which is quicker and less destructive to estimate than electivity indices. As a result, we demonstrate a useful technique for predicting herbivore diet selection.