Effects of fighting decisions on formation and structure of dominance hierarchies

The characteristics of complex social systems often cannot be predicted from exploring individual agonistic interactions in isolation. In a series of models, this study linked the decision-making processes that govern dyadic interactions with the emergent properties of hierarchical structures in social groups. Contrary to the intuitive expectation that resolution of ambiguities in fighting between closely matched opponents effectively promotes the formation of a social hierarchy, engagement in contests with opponents of dissimilar status lead to a faster emergence of hierarchical structures. The increased certainty of outcome in such asymmetric contests both reinforced the previously established dominance, and allowed for an indirect resolution of existing ambiguous relationships. High-return fights also resulted in a more rapid hierarchy formation, while escalating in fights decreased the costs of maintaining hierarchical relationships and increased their stability.     

Effects of fighting decisions on formation and structure of dominance hierarchies

The characteristics of complex social systems often cannot be predicted from exploring individual agonistic interactions in isolation. In a series of models, this study linked the decision-making processes that govern dyadic interactions with the emergent properties of hierarchical structures in social groups. Contrary to the intuitive expectation that resolution of ambiguities in fighting between closely matched opponents effectively promotes the formation of a social hierarchy, engagement in contests with opponents of dissimilar status lead to a faster emergence of hierarchical structures. The increased certainty of outcome in such asymmetric contests both reinforced the previously established dominance, and allowed for an indirect resolution of existing ambiguous relationships. High-return fights also resulted in a more rapid hierarchy formation, while escalating in fights decreased the costs of maintaining hierarchical relationships and increased their stability.     

Effects of fighting decisions on formation and structure of dominance hierarchies

The characteristics of complex social systems often cannot be predicted from exploring individual agonistic interactions in isolation. In a series of models, this study linked the decision-making processes that govern dyadic interactions with the emergent properties of hierarchical structures in social groups. Contrary to the intuitive expectation that resolution of ambiguities in fighting between closely matched opponents effectively promotes the formation of a social hierarchy, engagement in contests with opponents of dissimilar status lead to a faster emergence of hierarchical structures. The increased certainty of outcome in such asymmetric contests both reinforced the previously established dominance, and allowed for an indirect resolution of existing ambiguous relationships. High-return fights also resulted in a more rapid hierarchy formation, while escalating in fights decreased the costs of maintaining hierarchical relationships and increased their stability.     

Origins of behavioural variability: categorical and discriminative assessment in serial contests

Can social behaviour be probabilistic? Classical results in evolutionary game theory seem to suggest that recognizable asymmetries between individuals in the likelihood of winning fights (i.e. in resource holding power, RHP) rule out the stable probabilistic behaviour associated with mixed strategies. Here, using a variant of the hawk-dove game, I show that these mixed strategies can be common in asymmetric contests, because opponents of similar RHP benefit by ignoring discernable differences to form RHP equivalence categories. This process of categorizing (i.e. opting not to distinguish an opponent's discernably different RHP from one's own) can be adaptive when assessment is imprecise or expensive and mistakes can lead to dangerous combat. For large RHP differences between opponents, discriminating by acting on the discernable RHP difference is evolutionarily stable. For moderate RHP differences, typically neither categorizing nor discriminating is stable, but categorizing predominates if social interactions are frequent. More frequent interactions imply that the social status ultimately achieved is more important in accumulating fitness; this reduces the probability that a loser in combat will accept the submissive role without additional fighting, where this probability of ‘surrender’ expresses another mixed strategy. These patterns appear to fit the establishment of social relationships in some species for which combat is dangerous, and the analysis represents a step towards understanding the emergence of dominance hierarchies.     

Functions of fights in territory establishment

Fights are often observed when prospective territory owners settle in patches of vacant habitat, but the function of these fights in space acquisition is obscure. This study tests two hypotheses about the effect of fights on subsequent space use patterns: first, that settlers win space by winning fights and, second, that fights encourage the establishment of mutually exclusive home ranges between opponents (i.e., "fights make neighbors"). The behavior of juvenile Anolts aeneus lizards was recorded as they established territories in patches of habitat in the field. In support of the fights-make-neighbors hypothesis, opponents whose last aggressive interaction was a fight were six times more likely to have mutually exclusive home ranges at the end of the settlement period than were otherwise equivalent dyads whose last encounter was a chase. Contra the hypothesis that settlers win space by winning fights, most last fights ended in a draw, and there was no discernable relationship between the outcome of last fights and the subsequent space use of the contestants. These and previous analyses of settlement behavior in this species suggest that fights during the settlement period encourage the formation of symmetrical social and spatial relationships between neighboring settlers.     

Weaponry, color, and contest success in the jumping spider Lyssomanes viridis

Weaponry and color badges are commonly theorized to function as visual signals of aggressiveness or fighting ability. However, few studies have supported a signaling function of weaponry, and the role of color in invertebrate competitive interactions remains virtually unexplored. Jumping spiders (Salticidae) make excellent invertebrate models for studying weaponry and color because males of many species are colorful and possess exaggerated chelicerae, which are used as weapons in escalated contests. To determine whether color or weaponry might function as visual signals in male-male competitions, we investigated relationships between contest success, cheliceral length, and red coloration in Lyssomanes viridis. Males having longer chelicerae than their opponents were significantly more likely to win (p=0.0008). Males who won, despite being smaller than their opponents, had significantly less red chelicerae than their opponents (p=0.01). Male and female cheliceral length, as well as foreleg length, correlated tightly with body size. Cheliceral and foreleg length showed significantly stronger positive allometry in males than in females. We conclude that male chelicerae and forelegs are under strong positive selection for their use in physical fights and/or as visual signals of fighting ability.     

White-faced capuchin monkeys show triadic awareness in their choice of allies

The social intelligence hypothesis, which holds that social challenges have selected for increased intelligence and social skills, has been supported by evidence that, in catarrhine primates, individuals know about the characteristics of groupmates' social relationships. Evidence for such ‘triadic awareness’ has not been sought for platyrrhine primates, although two platyrrhine genera, capuchins, Cebus, and squirrel monkeys, Saimiri, are among the most highly encephalized primates. We examined patterns of coalitionary recruitment in wild white-faced capuchins, C. capucinus. Analyses have shown that more dominant individuals are more likely to join aggressive coalitions than low-rankers, and that individuals preferentially support those with whom they have stronger affiliative relationships. Data from 110 fights, analysed using simulation techniques that produced distributions of results expected under null hypotheses, revealed that contestants preferentially solicited prospective coalition partners that (1) were dominant to their opponents, and (2) had better social relationships (higher ratios of affiliative/cooperative interactions to agonistic interactions) with themselves than with their opponents. Further analyses showed that soliciting dominant partners could be explained by either of two simpler rules, ‘Solicit an ally that outranks yourself’ or ‘Solicit the highest-ranking available individual’. However, soliciting partners with better social relationships appears to indicate triadic awareness, because subjects did not preferentially solicit the nearby individual with whom they shared the highest-quality social relationship. Effects of relative relationship quality on coalition solicitation decisions were independent of effects of dominance rank.     

The smell of success and failure: the role of intrinsic and extrinsic chemical signals on the social behavior of crayfish

Animals commonly modify their behavior in the presence of aconspecific or in response to signals. This is particularlytrue in the context of aggressive exchanges, which animals useto form networks of social relationships and to communicatesocial status associated with those relationships. Althoughhierarchical structures are a widespread phenomenon that hasbeen studied extensively, the dynamic communication processes,specifically chemical communication in this review, is relativelyoverlooked. In particular, it is the exchange of informationduring agonistic interactions that mediates hierarchies and/oralters the outcomes of agonistic interactions. Given the theoreticalappeal of these interactions, and the evolutionary importanceand taxonomic diversity associated with social hierarchies,it is not surprising that the sensory mechanisms involved inthe formation and maintenance of hierarchical structures havereceived recent attention. In crayfish, dominance is thoughtto be largely determined by physical superiority, where encountersare largely dyadic and fighting behavior is highly stereotyped.However, recent evidence has shown that the outcome of dyadicencounters are dependent upon a number of factors other thanphysical size, that include the exchange of chemical informationduring encounters, previous social history, and the intrinsicneurochemical state of opponents. We have attempted to providea comprehensive analysis of the extrinsic chemical processes(previous history, sensory communication, etc.) and intrinsicchemical processes (neurochemical state) that produce and maintaindominance relations and social hierarchies in crayfish. We hopethat this review will bring together a global picture of theprocesses that determine a crayfish's social standing and howintrinsic and extrinsic chemicals have substantial effects onaggressive states and agonistic bouts.     

Changes in Heart Rate Associated with Contest Outcome in Agonistic Encounters in Lobsters

Agonistic contests between lobsters housed together in a confined space progress through encounters of increasing intensity until a dominance relationship is established. Once this relationship is established, losing animals continually retreat from the advances of winners.These encounters are likely to consume much energy in both winning and losing animals. Therefore, one might expect involvement of many physiological systems before, during and after fights. Here, we report effects of agonistic encounters on cardiac frequency in winning and losing adult lobsters involved in dyadic interactions.The results show that: (i) small but significant increases in heart rate are observed upon chemical detection of a conspecific; (ii) during agonistic interactions, further increases in heart rate are seen; and (iii) ultimate winners exhibit greater increases in heart rate lasting longer periods of time compared to ultimate losers. Heart rate in winners remains elevated for at least 15 min after the contests have ended and animals have been returned to their home tanks. Reduced effects are seen in second and third pairings between familiar opponents.The sustained changes in heart rate that we observe in winning lobsters may result from hormonal modulation of cardiac function related to the change in social status brought about by contest outcome.     

Contextual Factors Meditating Contests Between Male Chacma Baboons in Botswana: Effects of Food,Friends and Females

We examined aggressive displays among male chacma baboons (Papio ursinus) over a 23-mo period in the Okavango Delta of Botswana. High-ranking males were more likely than middle- or low-ranking males to participate in displays. Regardless of rank, all males were more likely to participate in chases or physical fights if their opponents rank was similar to their own. Most chases and fights, including those that led to injuries, were also between similarly-ranked males. The rate of both aggressive displays and approach-retreat interactions increased in the weeks before rank reversals, suggesting that rank challenges were preceded by a period when males assessed each others competitive ability and/or motivation. Aggressive displays between disparately-ranked opponents occurred most frequently in contests involving resources of high fitness value: the defense of meat, the defense of estrous females, and the protection of infants against infanticidal attacks. Silent displays were more likely to occur in these three contexts than were displays that occurred as part of more slowly escalating interactions, in which opponents first exchanged calls. Results suggest that competitive encounters among male baboons follow patterns predicted by evolutionary game theory.     

Influence of asymmetries in resource quality and age on aggression and dominance in black-chinned hummingbirds

According to current theory, dominance relationships may depend on asymmetries between adversaries in their expected net gain from contested resources. A prediction derived from this theory was confirmed in the field by experimentally creating asymmetries between neighbouring territorial hummingbirds with regard to their net energy gains from their territories, and then forcing them to fight for ownership of the combined resources of the two territories. When territory productivity was increased to levels well above the daily energy requirements of each contestant, energetic asymmetries no longer detectably influenced dominance, and the amounts of aggression per contest were lower. Contests in which an energetic asymmetry was partially offset by an age-related asymmetry involved more fights than contests in which opponents belonged to different age classes but controlled feeders of identical productivity. The predicted negative correlation between the magnitude of the asymmetry in net energy gain and agonistic investment in the contest did not occur.     

20-hydroxyecdysone causes increased aggressiveness in female American lobsters, Homarus americanus

Lobsters become transiently more aggressive before ecdysis. This aggressiveness accompanies an increase in hemolymph titers of 20-hydroxyecdysone (20-HE). Combats between intermolt female lobsters, injected with premolt levels of 20-HE, and larger, saline-injected opponents were videotaped. Aggressive, defensive, and avoidance behaviors were ranked according to aggressiveness in a Rank of Aggression hierarchy, which included opponent-directed and (nonopponent) redirected behaviors. Treated animals performed more and more highly aggressive behaviors than saline-injected controls. Opponents of treated animals performed fewer aggressive behaviors than saline-injected control opponents. Controls performed more defensive behaviors than treated animals, when redirected behaviors were considered. Differences in avoidance behaviors among the four types of combatants were not significant. The total aggressive content was the same in treated and control fights, but the interactions between combatants in the two fights were significantly different. Treated animals were equally as aggressive and defensive as their opponents; controls were relatively less aggressive and more defensive than their opponents. These results correlate with molt-cycle variations in behavior, 20-HE titers, and the effects of 20-HE and molt-differentiated hemolymph on the claw opener muscle. They suggest that 20-HE orchestrates intrinsic, cellular, and nuclear events that produce the molt-cycle transformations in agonistic behavior and aggressive state of lobsters.     

Aggression and RHP in the Northern Swordtail Fish, Xiphophorus cortezi: The Relationship Between Size and Contest Dynamics in Male–Male Competition

Current theory predicts that contest outcome, as well as decisions on whether to initiate a contest, escalate during a contest or retreat are decided by asymmetries in resource holding potential (RHP) and/or expected payoffs between contestants. In this investigation, dyadic contests were staged between male swordtail fish (Xiphophorus cortezi) where individuals were paired based on cumulative fight records and were ranked at the end of the trials in order to approximate RHP. Size was the only asymmetry that I did not attempt to control for and as a result, I was able to determine the relationships between size, contest initiation, escalation and outcome. Individuals changed their contest initiation strategy based on their size relative to that of their opponents, and contrary to predictions, the smaller of the two males in each contest was more likely to initiate the conflict than was the larger male. However, the larger of the two males was more likely to win and standard length proved to be a moderate predictor of an individual's final rank. Regardless of size, initiators fared poorly, winning only 31% of the contests. In instances where the smaller males won the contests, they were no more likely to initiate the encounter than was the larger male. However, when small males did win, fights lasted longer, suggesting that in some cases smaller males may be able to outlast their opponents.     

The role of individual differences and patterns of resolution in the formation of dominance orders in domestic hen triads

This research compares the role of initial individual characteristics to that of patterns of resolution in which successive dominance relationships are established during the formation of triads in the domestic hen. Combining weight and comb size with prior victory or defeat in the site of encounter, we created three levels of asymmetries of characteristics for triads of hens. The effects of these asymmetries were then examined on the resultant hierarchies and on the order of conflict resolution within triads under two different conditions of assembly. In one condition (simultaneous triad), the three hens were simultaneously introduced to each other and could thus freely choose their opponent. In the other condition (step-assembled triad), the hen predicted to occupy the highest rank was left on standby and introduced once the other two hens had settled dominance. This condition disrupts the normal process of hierarchy formation by imposing the first sequence of dominance settlement. We found that the structure of triadic hierarchies can be predicted from individual characteristics existing prior to hierarchy formation. No difference in the resultant structures were found between conditions of introduction, though different paths of conflict resolution were followed indicating that individual differences had a more determining role on the resultant hierarchies than patterns of resolution. In addition to demonstrating that individual differences determine resultant triadic structures, the present results also show that the same end structures can be reached by following resolution paths that are not necessarily of the Double Dominance and Double Subordinance types as prescribed by Chase's model. It is also found that in the simultaneous condition hens select each other to form pairs. Therefore, individuals do not meet at random but choose each other as opponents. The two hens predicted from individual differences to occupy the highest ranks first settle dominance, followed by settlement between the winner of the previous encounter and the bystander.     

Familiarity influences body darkening in territorial disputes between juvenile salmon

Escalated contests between animals are potentially costly because of increased energy expenditure and risk of predation or injury. Hence we would expect selection to favour any mechanism that avoids unnecessary prolonged fighting. One such means of avoiding escalated fights could be the use of information gained through individual recognition. Previous work has shown that a darkening of the body colour is closely associated with submission in contests between juvenile Atlantic salmon, Salmo salar, and it has been hypothesized that this may act as a visual signal to the opponent. We tested the hypothesis that body darkening is used to reduce the cost of contests between familiar fish such that losers darken more quickly when faced with familiar than unfamiliar opponents. In contests between unfamiliar fish, submissive darkening occurred after more escalated contests in which the loser incurred more aggression, whereas the opposite occurred when familiar fish were in conflict. In addition familiar fish either submitted quickly or engaged in protracted conflicts in which neither fish signalled submission, whereas in unfamiliar fish contests were of intermediate duration regardless of whether either fish darkened. We suggest that body darkening is used by familiar fish to signal submission to familiar dominants in order to avert a costly escalated fight, but familiarity can lead to escalation without submission if perceived competitive asymmetries are finely balanced. Copyright 2000 The Association for the Study of Animal Behaviour.     

Post-conflict Social Events in Wild Mountain Gorillas. II. Redirection,Side Direction,and Consolation

Inevitably, members of primate groups sometimes have aggressive contests with each other. Targets of aggression can engage in several types of interaction with third parties to ameliorate its adverse effects. They can redirect aggression, which may reduce the risk of further aggression from the initial opponents and reduce tension, or they can initiate affinitive interactions with third parties, in order to seek protection or reassurance. They can also receive reassuring acts from them (‘consolation’). Researchers have documented high levels of redirection in many primate species, but consolation is thus far known only in chimpanzees. Data on post-conflict social interactions between targets of aggression and third parties in two groups of wild mountain gorillas (Gorilla gorilla beringei) show that immatures and subordinate males, but not females, redirect aggression at high rates. Furthermore, immatures seek affinitive interactions with their mothers, and adult females seek them with adult males, at elevated rates. Affinitive interactions between females and males seem mainly to have a protective function and are associated with decreased risks of further aggression between female opponents. Redirection by females after female-female conflicts may be uncommon because targets commonly retaliate against aggressors. Males may offer females protection and consolation as services, and the results support the argument that such services are important when dominance relationships between females are often undecided and retaliation between opponents is common.     

Modelling dominance hierarchy formation as a multi-player game

Animals who live in groups need to divide available resources amongst themselves. This is often achieved by means of a dominance hierarchy, where dominant individuals obtain a larger share of the resources than subordinate individuals. This paper introduces a model of dominance hierarchy formation using a multi-player extension of the classical Hawk-Dove game. Animals play non-independent pairwise games in a Swiss tournament which pairs opponents against those which have performed equally well in the conflict so far, for a fixed number of rounds. Resources are divided according to the number of contests won. The model, and its emergent properties, are discussed in the context of experimental observations.     

Social inertia in white-throated sparrows results from recognition of opponents

Social inertia is a term for the stability of dominance relationships despite changes in the intrinsic dominating abilities of opponents. In a standard test for social inertia, low-ranking birds in an established hierarchy receive implants with testosterone (treated) and high-ranking birds receive empty implants (untreated). Social inertia occurs when the treated birds remain subordinate to untreated opponents in these groups, despite evidence that similarly treated birds dominate untreated strangers. In previous demonstrations of social inertia, however, treated and untreated birds were returned to their original aviaries and tested with familiar opponents, and thus the effects of familiarity with the location and those of familiarity with opponents were not separated. To address this issue, we investigated social inertia in 16 groups of white-throated sparrows Zonotrichia albicollis. When low-ranking treated birds were placed in new aviaries with familiar high-ranking, untreated opponents (treatment S, same opponents), dominance relationships showed social inertia. When such birds were placed in new aviaries with unfamiliar opponents (treatment N, new opponents), testosterone influenced dominance. When groups of high-ranking, untreated birds acquainted with each other were placed with unfamiliar treated opponents (treatment G, grouped dominants), 'coat-tail' effects (dominance by association with high-ranking individuals) sometimes outweighed the effects of testosterone. Social inertia in this species is thus a result of familiarity with opponents, rather than familiarity with locations of encounters. Measurements of aggressive tendencies confirmed a previous report that social inertia suppresses activation of aggression by testosterone. White-throated sparrows can thus recognize their opponents, and this ability affects the expression of both dominance and aggression. Copyright 1999 The Association for the Study of Animal Behaviour.     

The assessment game in sand fiddler crab contests for breeding burrows

The single enlarged claw of male sand fiddler crabs, Uca pugilator, is used in contests for control of breeding burrows. The larger of the two contestants has the larger claw and usually wins. Males use one or more of 10 agonistic elements that vary in intensity from a no-contact extension of the claw to the flip of an opponent. We used the sequence of elements employed and the duration of unstaged, naturally occurring contests in a South Carolina salt marsh to evaluate three models of extended contests: (1) energetic war of attrition, (2) cumulative assessment and (3) sequential assessment. Contests usually began with elements of low action intensity and often proceeded to elements of high intensity. Elements of higher intensity were correlated with both contest duration and the number of contest elements. Contest duration increased as opponents became more evenly matched in size, a result consistent with both cumulative and sequential assessment models. Variation in duration increased as the relative sizes of opponents increased, also in accordance with sequential assessment. The absolute size of the smaller contestant had no effect on contest duration, in contrast to predictions based on cumulative assessment or energetic war of attrition models. Contestants that lost a fight were more likely to engage immediately in another fight without loss of contest intensity, if their previous fight had been long and intense. This result is inconsistent with contests of endurance, such as the energetic war of attrition or the cumulative assessment game, but it is consistent with the ritualized display of strength and fighting skill. Thus, sequential assessment appears to best explain ritualized fiddler crab contests. Cumulative assessment, however, may be the appropriate model for extended, nonritualized, all-out fights. Cumulative assessment may also explain the tenure of individuals on breeding grounds where multiple engagements are likely to test endurance and tolerance to damage over a period of days. Copyright 2003 Published by Elsevier Science Ltd on behalf of The Association for the Study of Animal Behaviour.      

Reconciliation in the Spotted Hyena (Crocuta crocuta)

Theory predicts that it should often be in the best interests of gregarious animals to repair social bonds damaged by within-group conflict. Indeed, reconciliation in many primates takes the form of affiliative behavior occurring shortly after a conflict. Here we inquired whether reconciliation also occurs among spotted hyenas ( Crocuta crocuta ), gregarious carnivores whose social lives share much in common with those of cercopithecine primates. In a large group of free-living hyenas in Kenya, we used focal animal observations to monitor rates at which various affiliative behaviors occurred before and after dyadic aggressive interactions. An affiliative behavior was identified as having a conciliatory function only if it occurred more frequently after than before fights, and if it was also associated with reduced rates of aggression between former opponents during the post-conflict interval. Of all affiliative behaviors monitored, only two types satisfied both these criteria: greeting behavior and non-aggressive approach. Over 72% of conciliatory behaviors occurred within the first five minutes after a fight, and hyenas reconciled after 14.6% of 698 fights. Mean conciliatory tendency (CT) for individual hyenas was 11.3%. Hyenas exhibited higher CTs when they were recipients (victims) of aggression than when they were aggressors, and they showed higher CTs in interactions with non-kin than with kin. Conciliatory tendencies did not vary with age–sex classes of opponents or with rank distance between opponents . Conciliatory tendency in spotted hyenas fell at the low end of the CT range observed among primates.