Development of Badges of Status in Captive Male House Sparrows (Passer domesticus) in Relation to the Relative Ornamentation of Flock‐Mates

The development of quality signals is expected to be largely influenced by environmental conditions among which the social environment could be an important factor, particularly in gregarious animals. Here, we investigated the role of the presumed social environment on the development of badge size in moulting male House Sparrows (Passer domesticus) over a 3‐yr time span. House Sparrow badges are testosterone‐dependent ornaments that are typically argued to be ‘badges of status’ –that is, signals of competitive ability and aggression. Immediately prior to moult, we created several groups of males, either with similarly sized badges or with badges of different sizes, and we measured each male's badge size before and after moult. The former groups were presumed to have a less clear social hierarchy than the latter groups because of group members advertising relatively similar competitive ability (and thus were expected to more frequently challenge one another in order to establish dominance). Conversely, the latter groups were presumed to have relatively higher social stability because birds would less frequently encounter rivals advertising similar agonistic threat. We predicted in groups of birds with similarly sized badges, both (1) larger changes (i.e. increases or decreases) in badge sizes and (2) higher increases in between‐individual variation within groups. The first prediction was supported in one of the 3 yrs, but there was no support for the second prediction in any year. We conclude that social environment has an effect on the development of badges of status, but that it is highly contingent on the specific nature and timing of group interactions.     

Natural and sexual selection on a plumage signal of status and on morphology in house sparrows,Passer domesticus

Temporal patterns of natural and sexual selection on male badge size and body traits were studied in a population of house sparrows, Passer domesticus. Badge size was a heritable trait as revealed by a significant father-son regression. Survival during autumn dispersal and winter was not related to badge size or body traits in yearling male house sparrows. Badges that signal dominance status were affected positively by directional selection for mating. Adult male house sparrows suffered an opposing selection pressure on badge size during autumn. Contrary to males, female house sparrows did not experience significant directional or stabilizing selection on any body trait. Directional sexual selection on male badge size due to female choice moves male sparrows away from their survival optimum. Opposing directional natural selection on badge size due to autumn mortality caused by predation maintains a stable badge size.     

Social Control and Physiological Cost of Cheating in Status Signalling Male House Sparrows (Passer domesticus)

Flock-forming passerines often use plumage characteristics to signal their social dominance. While the benefits to signal dominance seem obvious, costs associated with status signalling are ambiguous. The social control hypothesis predicts that individuals of high social status – with large badges – are involved in more social interactions with individuals of similar badge size. Cheaters are therefore exposed to increased risk of fighting with high quality individuals and the costs associated with enhanced fights with dominant males are supposed to outweigh the benefits of cheating. We tested the social control hypothesis in male house sparrows ( Passer domesticus ), by observing social interactions in captive flocks and determining dominance relationships. Two low status individuals within each flock had the size of their badge experimentally increased and the interactions involving experimental and control birds were recorded. We also assessed the potential physiological cost of cheating in terms of enhanced levels of the stress hormone, corticosterone. Dominance was significantly positively correlated with badge size, but not with other morphological traits. We found little support for the social control hypothesis. Birds did not have significantly more interactions with individuals of similar badge size, before the manipulation. Similarly, after the experimental increase in badge size, experimental birds did not tend to have more encounters with large-badged males. Experimental birds with enlarged badges won more fights compared with prior to the manipulation, suggesting that badge size is used as a signal of social dominance even in small and stable flocks. Finally, corticosterone levels in the blood did not increase significantly after the manipulation of badge size, suggesting that there is no measurable cost, resulting from stress, in cheaters.     

Benefits and Costs of Dominance in the Angelfish Centropyge bicolor

Social groups are often structured by dominance hierarchies in which subordinates consistently defer to dominants. High‐ranking individuals benefit by gaining inequitable access to resources, and often achieve higher reproductive success; but may also suffer costs associated with maintaining dominance. We used a large‐scale field study to investigate the benefits and costs of dominance in the angelfish Centropyge bicolor, a sequential hermaphrodite. Each haremic group contains a single linear body size‐based hierarchy with the male being most dominant, followed by several females in descending size order. Compared to their subordinate females, dominant males clearly benefited from disproportionately high spawning frequencies, but bore costs in lower foraging rates and greater aggressive defence of their large territories. Within the female hierarchy, more dominant individuals benefited from higher spawning frequencies and larger home ranges, but displayed neither higher foraging rates nor spawn order priority. However, dominance in females was also linked to aggressiveness, particularly towards immediate subordinates, suggesting that females were using energetically costly aggression to maintain their high rank. We further showed by experimentally removing dominant females that the linear hierarchy was also a social queue, with subordinates growing to inherit higher rank with its attendant benefits and costs when dominants disappeared. We suggest that in C. bicolor, the primary benefit of high rank is increased reproductive success in terms of current spawning frequency and the prospect of inheriting the male position in the near future, which may be traded off against the cost of aggressively defending rank and territory.     

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.     

Does Male Badge Size Signal Status in Small Island Populations of House Sparrows,Passer domesticus?

The relationship between dominance and throat badge size, body size, and body condition was examined in three small island populations of house sparrows, Passer domesticus, on the coast of northern Norway. Individual dominance rank in males was determined by observation of agonistic interactions in an artificial observation cage (one replicate in Dec. and one in Mar. in two populations and one replicate in Mar. in the third population). Previous studies had indicated that badge size signals status in house sparrows. In this study, because of the small population sizes and stability in their membership, we expected that prior information about fighting ability would reduce the importance of badge size for the outcome of conflicts in favour of traits most directly associated with fighting ability. In two of five replicates, significant relationships were found between components of body size and dominance rank, but in opposite directions. The relationship between badge size and dominance rank was consistently positive, but non-significant. In a pooled sample of the three independent replicates, badge size was the best predictor of dominance, and no other variable explained any variance in dominance. However, the coefficient of determination was low and linearity of dominance hierarchies was poor. We therefore suggest that individual recognition may influence the dominance relationships.     

Multiple Cues in Status Signalling: The Role of Wingbars in Aggressive Interactions of Male House Sparrows

During aggressive interactions, animals may signal their competitive ability by various ornaments referred to as badges of status. The use of a single badge predicting dominance rank occurs in many vertebrate species. However, animals often display multiple ornaments that may convey information about either different or the same aspects of the signaller's quality, or alternatively, may serve as signal amplifiers. We observed the fighting behaviour of male house sparrows in two captive flocks to investigate whether they may use multiple cues in status signalling during aggressive interactions. Beside the status‐signalling bib, male sparrows possess a conspicuous white wingbar that they often display upon aggressive encounters. We tested whether bib size and the wingbar's conspicuousness (i.e. its achromatic contrast with the neighbouring dark feathers) or its area predicted success in various aspects of fighting. We found that bib size strongly predicted overall fighting success (i.e. proportion of fights won) and defence success (i.e. proportion of successful defences out of all attacks received). Wingbar conspicuousness was positively related to defence success after controlling for the effect of bib size in multivariate analyses. Furthermore, displaying the wings also tended to improve the birds’ success in defence but not in attack. Wingbar area was unrelated to any measured aspect of fighting ability. We suggest that bib size and wingbar conspicuousness may convey multiple messages on fighting abilities, specifically on overall aggressiveness and defending potential, respectively. Alternatively, wingbars may serve as amplifiers for the wing displays of aggressive motivation. Thus, male sparrows may use multiple cues in assessing the competitive ability of opponents during social interactions.     

Bimorphism in Male Verreaux’s Sifaka in the Kirindy Forest of Madagascar

Male primates in species with pronounced secondary sexual adornments can exhibit reversible or irreversible bimorphism, i.e., striking variation in the degree to which males express the adornments. Verreaux’s sifaka (Propithecus verreauxi verreauxi) use scent marking as a form of communication and exhibit sex differences in scent glands. Some males exhibit a pronounced brown staining around their sternal gland, whereas others do not. We studied morphological and behavioral characteristics of males in 6 social groups in Kirindy Forest, Madagascar, from November 2000 to March 2002 to evaluate the hypotheses that the bimorphism in male sifaka chest status represents alternative mating tactics and is a badge of status. Males are clearly divided into 2 categories: clean and stained chests, with rare, but informative, intermediate males. The chest staining probably results from the males scent marking with their sternal glands, because stained-chested males scent marked significantly more often than clean-chested males. Though sample sizes are small, chest status did not appear to depend on body size. Chest status is reversible and related to dominance rank. In each group, only 1 male, the dominant, was stained-chested, whereas all other (subordinate) males were clean-chested. These findings suggest that stained chests are visual and olfactory signals of dominance rank and that clean chests signal lack of competitive intent. Thus, this bimorphism may reflect alternative mating tactics used by males to maximize their reproductive success based upon their social environment.     

Information content of male agonistic displays in the territorial tawny dragon (Ctenophorus decresii)

Two potential signals used during male–male agonistic encounters were examined for signal content in the territorial agamid lizard Ctenophorus decresii, or tawny dragon. Males have black chest patches, which are apparent when they posture during agonistic encounters. Patches are not condition or size dependent. The area of the patches is positively associated with levels of aggression and likelihood of winning a fight. The patch thus functions as a badge of status indicating male aggression. The complex dynamic displays given by males contain information on male endurance and size. The number of push-ups given during a display reflects the aggressiveness of an animal. There was no relationship between patch size and endurance. There is some overlap in the content of the two signals, both contain information on aggressiveness, suggesting that they may function as back-up signals. The multiple-message hypothesis is not ruled out as endurance and size are only related to the dynamic displays. However, it is not clear that endurance is an important determinant of contest outcomes in this species, and so it is not certain that the receiver uses this information.     

Sex reversal in pairs of Lythrypnus dalli: behavioral and morphological changes

In Lythrypnus dalli, the bluebanded goby, reproductive success is primarily determined by functional sex, and functional sex is determined largely by rank in the dominance hierarchy. In most natural social groups of L. dalli, one male is at the apex of the hierarchy, and 1 to 7 females are lower in rank. When a male exits the group, a female ascends to the top of the hierarchy and becomes a male. We have examined this process in a simplified environment--a pair of females--that allows us to identify behavior associated with the formation of a dominance relationship and any other phenotypic changes associated with dominance, sex change or both. We found that pairs of L. dalli females quickly and readily form stable dominance relationships, with the dominant fish changing sex into a male. This dominant animal also rapidly increases in body size and length of its dorsal fin. In summary, dominant L. dalli females change sex in this simplified environment, providing excellent opportunities to examine the early behavioral and morphological changes associated with dominance and sex change.     

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.     

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.     

Trade‐offs in primate grooming reciprocation: testing behavioural flexibility and correlated evolution

Primates may trade altruistic behaviours, such as grooming, either for itself or for different rank‐related benefits, such as tolerance or agonistic support. Ecological conditions are expected to affect competition and thus the steepness of dominance hierarchies. This, in turn, may influence the value of the different currencies that primates exchange. Thus, it can be hypothesized that, as the dominance hierarchy becomes steeper, more grooming is directed up the hierarchy (in exchange for tolerance or agonistic support) and less grooming is exchanged for other grooming. We assembled a large database of within‐group grooming distribution in primates (38 social groups belonging to 16 species and eight genera) and tested these hypotheses both within species (i.e. comparing different groups of the same species) and between species (using comparative methods that control for phylogenetic relatedness). We found within‐species evidence that steeper dominance hierarchies were associated with more grooming being directed up the hierarchy, and that a trade‐off occurred between the tendency to groom up the hierarchy and the degree of grooming reciprocation (although, in some analyses, only a nonsignificant trend was observed). By contrast, phylogenetically controlled comparisons between species did not reveal evidence of correlated evolution between the steepness of the dominance hierarchy, the tendency to direct grooming up the hierarchy, and the degree of grooming reciprocation. © 2008 The Linnean Society of London, Biological Journal of the Linnean Society, 2008, 95 , 439–446.     

Group Size in Animal Societies: The Potential Role of Social and Ecological Limitations in the Group‐Living Fish,Paragobiodon xanthosomus

The number of group members in an animal society can have a major influence on group members’ life history, survival, and reproductive success. Identifying the factors that limit group size is therefore fundamental for a complete understanding of social behavior. Here, I examined the relationships between resource availability, social conflict, and group size in the coral‐dwelling fish, Paragobiogon xanthosomus (Gobiidae). The size of the largest (breeding) female and the minimum size difference required for hierarchy stability strongly but not perfectly predicted maximum group size, suggesting that social conflicts and hierarchy structure set the upper limit on group size. Deviations in group size around the predicted maximum were explained by variation in average body size ratios but not by variation in coral size, suggesting that coral size does not directly influence group size. In contrast, coral size was a significant predictor of body size ratios, and possible explanations for this relationship are discussed. Group size may be limited by the social conflicts that characterize size‐based dominance hierarchies. Ecological factors, namely coral size, may in turn play an indirect role via an effect on body size ratios.     

Affiliative relationships among Sorraia mares: influence of age, dominance, kinship and reproductive state

The influence of individual factors on dominance rank and the relationship between rank distance and patterns of aggression predicted by models of evolutionarily stable strategies (ESS) of animal conflict were investigated in a managed bachelor group of Sorraia horses, Equus caballus. The group was composed of four to six stallions 3- to 12-years-old during the study period. The dominance hierarchy was significantly linear and rank was not related to age, weight, height or aggressiveness. Frequency and intensity of agonistic interactions were low, but higher-ranking stallions did not receive lower aggressiveness than lower-ranking stallions. There was some evidence that dominance relationships were more contested among close-ranking stallions, as predicted. Agonistic-related interactions among close-ranking stallions served similar functions to those among distant-ranking stallions, but the latter interacted more frequently than expected for access to resting sites and/or resting partners. Therefore, we found some evidence that agonistic-related interactions among distant-ranking stallions play a larger role in providing access to valuable and defendable resources than those among close-ranking stallions. Nevertheless, the fact that space to escape from aggression was limited and breeding access was independent from dominance rank may have reduced the benefits relative to costs of aggression and therefore limited the occurrence of contests over dominance and resources.     

Dominance, Status Signals and Coloration in Male Mandrills (Mandrillus sphinx)

Where individuals contest access to a resource, escalated physical fighting presents a risk to all involved. The requirement for mechanisms of conflict management has led to the evolution of a variety of decision rules and signals that act to reduce the frequency of aggression during competitive encounters. We examined strategies of conflict management in male mandrills (Mandrillus sphinx) living in two semi‐free‐ranging groups in Gabon. Adult male mandrills are large (31 kg), with long canines, making the costs of conflict potentially very high. We found that males formed dominance hierarchies, but that male–male relationships were characterized by avoidance, appeasement and ignoring. Fights were rare, but could result in death. Examination of the relationship between dominance and signaling showed that males use facial and gestural signals to communicate dominance and subordinance, avoiding escalated conflict. Male mandrills also possess rank‐dependent red coloration on the face, rump and genitalia, and we examined the hypothesis that this coloration acts as a ‘badge of status’, communicating male fighting ability to other males. If this is the case, then similarity in color should lead to higher dyadic rates of aggression, while males that differ markedly should resolve encounters quickly, with the paler individual retreating. Indeed, appeasement (the ‘grin’ display), threats, fights and tense ‘stand‐off’ encounters were significantly more frequent between similarly colored males, while clear submission was more frequent where color differences were large. We conclude that male mandrills employ both formal behavioral indicators of dominance and of subordination, and may also use relative brightness of red coloration to facilitate the assessment of individual differences in fighting ability, thereby regulating the degree of costly, escalated conflict between well‐armed males.     

Social relationships and social structure of the mound-building mouse (<Emphasis Type="Italic">Mus spicilegus</Emphasis>) in intraspecific cage groups

The social relationships of Mus spicilegus were studied in intraspecific cage groups for a month during the reproductive period to clarify the role of intraspecific aggressiveness in shaping social structure and in reproduction. The mound-building mice were captured from a wild population in northern Bulgaria. A total of 13 groups, 5 male, 4 female, and 4 male–female groups, were tested. Each group consisted of five unfamiliar adults. The experiments were carried out in 100×100-cm glass cages. The patterns of behaviour and the dominant–subordinate structure within each group were investigated. A dominance index for every individual in the group was calculated. During the period of establishment of the hierarchical structure in the groups a high level of aggressiveness was registered among individuals. About half of the subdominants and subordinates were found dead, with or without obvious marks of injury. Male–male interactions were more aggressive compared to these of females. Dominant–subordinate relationships between individuals of the same sex were observed in the mixed groups. A link between dominance rank and reproduction was established. The dominant male and the dominant female formed a pair and started to breed. Based on these findings it can be assumed that the aggressiveness of M. spicilegus is an adaptation to its life in the fields, where dominance is likely to be an essential factor in individual reproductive success. It is likely that the successful rearing of offspring in natural populations, which are exposed to strong biotic and abiotic impact, necessitates the existence of monogamous relationships. Communicated by P. Gama Mota     

迁地保育林麝的侵犯性、等级序位及相互关系

侵犯性是动物个性的重要维度之一,体现了其主动挑衅和攻击其他个体的倾向。动物的侵犯性与其社会结构及等级序位存在紧密关系。本研究于2018年6月1日至7月31日对四川马尔康林麝繁育场的圈养林麝(Moschus berezovskii)进行了行为取样,计算林麝个体的侵犯性个性(侵犯性指数)和等级序位指数,分析圈养林麝的侵犯性和等级序位格局、影响因素及相互关系。结果表明,圈养林麝的侵犯性在性别间存在显著差异,雄性林麝的侵犯性(0.45±0.09,n=22)显著高于雌性(0.22±0.06,n=30)(P <0.05),年龄和驯养密度对其侵犯性的效应均不显著(P> 0.05),说明林麝侵犯性的刚性较强;圈养林麝等级序位的性别间差异不显著(P> 0.05),亚成体麝与成体麝的等级序位差异也未达显著水平(P> 0.05),原因在于麝场的建群未区分年龄组;圈养林麝个体的侵犯性与其等级序位显著正相关,林麝个体的侵犯性越大,其等级序位越高(r=0.73,P <0.05),推测这与社群的序位等级构建和资源竞争有关。     

Status signalling, metabolic rate and body mass in the siskin: the cost of being a subordinate

The higher metabolic rate of dominant individuals, found in different species, has been interpreted as the cost that prevents subordinates from cheating by adopting large badges of status. However, an alternative prediction for status-signalling species, in which subordinates may recognize dominants, is that subordinates have the higher metabolic rate because of the greater stress of locating and actively avoiding aggressive interactions with them. In this study, the size of the black bib of the siskin, Carduelis spinus, which is a badge of dominance, was negatively correlated with metabolic rate in daylight, even when controlling for the bird's activity level in the respirometer chamber and its body mass. The size of the black bib, however, was not correlated with metabolic rate in darkness. This suggests that the difference between dominance classes is not related to intrinsic physiological differences, but that subordinates are more susceptible to stressful conditions. When controlling for metabolic rate, a positive correlation appeared between dominance status and body mass. This stresses the importance of knowing the effects of social status on energy requirements for understanding the relationship between body mass and dominance. We conclude that maintaining a high social status may be more stressful to subordinates than to dominant birds. Copyright 2000 The Association for the Study of Animal Behaviour.     

Memory of Social Partners in Hermit Crab Dominance

We investigated the possibility that invertebrates recognize conspecific individuals by studying dominance relationships in the long‐clawed hermit crab, Pagurus longicarpus. We conducted three sets of laboratory experiments to define the time limits for acquiring and maintaining memory of an individual opponent. The results reveal two characteristics that make individual recognition in this species different from standard associative learning tasks. Firstly, crabs do not require training over many repeated trials; rather, they show evidence of recognition after a single 30‐min exposure to a stimulus animal. Secondly, memory lasts for up to 4 d of isolation without reinforcement. A third interesting feature of individual recognition in this species is that familiar opponents are recognized even before the formation of a stable hierarchical rank. That is, recognition seems to be relatively independent of repeated wins (rewards) or losses (punishments) in a dominance hierarchy. The experimental protocol allowed us to show that this species is able to classify conspecifics into two ‘heterogeneous subgroups’, i.e. familiar vs. unfamiliar individuals, but not to discriminate one individual of a group from every other conspecific from ‘a unique set of cues defining that individual’. In other words, we demonstrated a ‘binary’– and not a ‘true’– individual recognition. However, 1 d of interactions with different crabs did not erase the memory of a former rival, suggesting that P. longicarpus uses a system of social partner discrimination more refined than previously shown.