A model of coalition formation in animals

A simple three-player model is presented for the evolution of coalitions. The model demonstrates that, under certain conditions, ''winner'' and ''loser'' effects both favour coalition formation. Winner effects are defined as an increased probability of winning at time T + 1, given a victory at time T, whereas loser effects entail an increased probability of losing at time T + 1, given a loss at time T. Increasing the strength of loser effects or winner effects, or the strength of an individual''s position in the hierarchy, makes coalition formation in general more likely, whereas increasing the costs of giving aid does the opposite. The model does not assume any form of reciprocity, but rather examines whether some form of reciprocity or pseudoreciprocity emerges from the model itself. When either winner or loser effects exist, reciprocal coalition formation (e.g. i helps j against k, and j helps i against k) between β (second-ranked individual) and α (highest-ranked individual) or between α and γ (lowest-ranked individual) was possible, but reciprocal aid-giving between γ and β was never favoured. Thus, we have the counterintuitive result that although a coalition between the two lowest members of a hierarchy against the dominant individual is possible (as selection may favour γ aiding β against α), such a coalition is not predicted to be reciprocal in kind. Interpopulational comparisons examining winner/loser effects and coalition formation would allow for a test of many of the model''s most basic predictions. Unfortunately, most work on coalitions has been undertaken in primates, whereas work on winner and loser effects has focused on rodents, and more recently, in fish and birds. Hopefully, the model presented here will spur future work that will look at all of these factors simultaneously in many taxa.     

Winner and loser effects and the structure of dominance hierarchies

In the literature on dominance hierarchies, "winner" and "loser"effects usually are denned as an increased probability of winningat time T, bated on victories at time T-l, T-2, etc, and anincreased probability of losing at time T, based on losing atT-1, T-2, etc., respectively. Despite some early theoreticalwork on winner and loser effects, these factors and how theyaffect the structure of dominance hierarchies have not beenexamined in detail. I developed a computer simulation to examinewinner and loser effects when such effects are independent ofone another (as well as when they interact) and when combatantsassess each other's resource-holding power. When winner effectsalone were important, a hierarchy in which all individuals heldan unambiguous rank was found. When only loser effects wereimportant, a dear alpha individual always emerged, but the rankof others in the group was often unclear because of the scarcityof aggressive interactions. Increasing winner effects for agiven value of the loser effect increase the number of individualswith unambiguous positions in a hierarchy and the converse istrue for increasing the value of the loser effect for a givenwinner effect Although winner and loser effects have been documentedin a number of species, no study has documented both winnerand loser effects (using some controlled, pairwise testing system)and the detailed nature of behavioral interactions when individualsare in groups. I hope the results of this model will spur suchstudies in the future.     

Coalition formation in animals and the nature of winner and loser effects

Coalition formation has been documented in a diverse array of taxa, yet there has been little formal analysis of polyadic interactions such as coalitions. Here, we develop an optimality model which examines the role of winner and loser effects in shaping coalition formation. We demonstrate that the predicted patterns of alliances are strongly dependent on the way in which winner and loser effects change with contestant strength. When winner and loser effects decrease with the resource-holding power (RHP) of the combatants, coalitions will be favoured between the strongest members of a group, but not between the weakest. If, in contrast, winner and loser effects increase with RHP, exactly the opposite predictions emerge. All other things being equal, intervention is more likely to prove worthwhile when the beneficiary of the aid is weaker (and its opponent is stronger), because the beneficiary is then less likely to win without help. Consequently, intervention is more probable when the impact of victory on the subsequent performance of a combatant increases with that individual's strength because this selects for intervention in favour of weaker combatants. The published literature on hierarchy formation does not reveal how winner and loser effects actually change with contestant strength and we therefore hope that our model will spur others to collect such data; in this light we suggest an experiment which will help to elucidate the nature of winner and loser effects and their impact on coalition formation in animals.     

Persistence of Winner and Loser Effects Depends on the Behaviour Measured

Recent contest experience can influence an individual’s behaviour in subsequent contests. When the probability of winning a subsequent contest is used to quantify experience effects, a loser effect usually lasts longer than a winner effect. This conclusion, however, may be caused by this probability understating the persistence of the influence of a winning experience on contest decisions. Using Kryptolebias marmoratus, a mangrove killifish, as the study organism, we investigated whether different conclusions about the relative persistence of winning and losing experiences would be reached when different aspects of contest behaviour (probability of initiating attacks, probability of winning non‐escalated and escalated contests, escalation rate and contest duration) were measured. The results indicated that the apparent persistence of the effect of winning or losing experiences varied with the behaviour studied. When the likelihood to initiate attacks was used, no winner effect was detected while the loser effect lasted for <1 d. When escalation rate was used, the winner effect lasted for 2–4 d, while the loser effect lasted for 1–2 d. When the probability of winning non‐escalated contests was used, the winner effect was detectable for <1 d, while the loser effect lasted for 2–4 d. And, when contest duration was used, the winner effect was detectable for 2–4 d, but no loser effect was detectable. These results show that (1) the probability of winning a subsequent contest understated the persistence of the influence of a winning experience on the fish’s contest decisions, (2) the measures most effective at detecting winner effects are different from those most effective at detecting loser effects and (3) in K. marmoratus, both effects can be detected 2 d after the completion of experience training but both dissipate in 4 d.     

Why do winners keep winning? Androgen mediation of winner but not loser effects in cichlid fish

Animal conflicts are influenced by social experience such that a previous winning experience increases the probability of winning the next agonistic interaction, whereas a previous losing experience has the opposite effect. Since androgens respond to social interactions, increasing in winners and decreasing in losers, we hypothesized that socially induced transient changes in androgen levels could be a causal mediator of winner/loser effects. To test this hypothesis, we staged fights between dyads of size-matched males of the Mozambique tilapia (Oreochromis mossambicus). After the first contest, winners were treated with the anti-androgen cyproterone acetate and losers were supplemented with 11-ketotestosterone. Two hours after the end of the first fight, two contests were staged simultaneously between the winner of the first fight and a naive male and between the loser of first fight and another naive male. The majority (88%) of control winners also won the second interaction, whereas the majority of control losers (87%) lost their second fight, thus confirming the presence of winner/loser effects in this species. As predicted, the success of anti-androgen-treated winners in the second fight decreased significantly to chance levels (44%), but the success of androgenized losers (19%) did not show a significant increase. In summary, the treatment with anti-androgen blocks the winner effect, whereas androgen administration fails to reverse the loser effect, suggesting an involvement of androgens on the winner but not on the loser effect.     

Recognition of Dominance in the Big-Clawed Snapping Shrimp ( Alpheus heterochaelis Say 1818) Part II: Analysis of Signal Modality

Alpheus heterochaelis is able to recognise the dominance status of an opponent (see Part I of this series). A former loser does not fight against a former winner but rather escapes immediately after a contact. However, if a former loser meets an inexperienced opponent, the loser fights against it. Here we investigated the signal used for dominance recognition. Two groups of snapping shrimp that had lost a fight on Day 1, intact animals, and shrimp with cut lateral antennular filaments (i.e. without chemosensory aesthetascs), fought against the same winner on Day 2. Intact losers showed escape behaviour, while losers without aesthetascs showed almost the same aggressive behaviour as on Day 1. The main signal in dominance recognition is therefore a chemical one, possibly the urine or a substance carried by it. The main receptor organs for this signal are the lateral filaments of the antennules carrying the aesthetascs.     

Winner and loser effects in human competitions. Evidence from equally matched tennis players

Animals winning an agonistic encounter are more likely to win their next encounter while losers are less likely, even when controlling for motivation and physical size. Do these winner and loser effects exist in human competitions? Drawing on a large database of professional tennis matches, we were able to control for players' ability and thereby test for winner and loser effects. We narrowed the database to matches between players who on average did not differ significantly in rank, and further to matches in which the first set was fought to a long tie-break. These closely fought matches present a natural experiment because players are assigned to treatment conditions – winning or losing a set – despite similar ability and performance. We found that among men, the winner of a closely fought tie-break had an approximate 60% chance of winning the second set, the loser a 40% chance. These effects did not exist among women, a finding consistent with the hypothesis that androgens mediate winner and loser effects. Our results may help in the design of competitions in sport as well as in work environments, where it may prove useful to either encourage winner effects or to attenuate their occasional adverse consequences.     

Recognition of Dominance in the Big-Clawed Snapping Shrimp (Alpheus heterochaelis Say 1818) Part II: Analysis of Signal Modality

Alpheus heterochaelis is able to recognise the dominance status of an opponent (see Part I of this series). A former loser does not fight against a former winner but rather escapes immediately after a contact. However, if a former loser meets an inexperienced opponent, the loser fights against it. Here we investigated the signal used for dominance recognition. Two groups of snapping shrimp that had lost a fight on Day 1, intact animals, and shrimp with cut lateral antennular filaments (i.e. without chemosensory aesthetascs), fought against the same winner on Day 2. Intact losers showed escape behaviour, while losers without aesthetascs showed almost the same aggressive behaviour as on Day 1. The main signal in dominance recognition is therefore a chemical one, possibly the urine or a substance carried by it. The main receptor organs for this signal are the lateral filaments of the antennules carrying the aesthetascs.     

Individual recognition, dominance hierarchies and winner and loser effects

Winner and loser effects are defined as an increased probability of winning an aggressive interaction at time T, based on victories at time T-1, T-2, etc., and an increased probability of losing at time T, based on losses at time T-1, T-2, etc., respectively. Prior theoretical work on dominance hierarchy formation has demonstrated that when players are not capable of individual recognition, loser effects always produce a clear top-ranked (alpha) individual, but all other ranks in a group remain unclear; whereas winner effects always produce strict linear hierarchies in which the rank of each individual is clear. Paradoxically, however, when individual recognition--a phenomenon long thought to stabilize hierarchies--is possible, winner and loser effects have no impact on the probability of forming strict linear hierarchies.     

Opponent familiarity and contest experience jointly influence contest decisions in <Emphasis Type="Italic">Kryptolebias marmoratus</Emphasis>

Introduction

Individual recognition and winner/loser effects both play important roles in animal contests, but how their influences are integrated to affect an individual’s contest decisions in combination remains unclear. Individual recognition provides an animal with relatively precise information about its ability to defeat conspecifics that it has fought previously. Winner/loser effects, conversely, rely on sampling information about how an animal’s ability to win compares with those of others in the population. The less precise information causing winner/loser effects should therefore be more useful to an individual facing an unfamiliar opponent. In this study, we used Kryptolebias marmoratus, a hermaphroditic mangrove killifish, to test whether winner/loser effects do depend on opponent familiarity. In addition, as previous studies have shown that subordinates that behave aggressively sometimes suffer post-retreat retaliation from contest winners, we also explored this aspect of contest interaction in K. marmoratus.

Results

In the early stages of a contest, subordinates facing an unfamiliar dominant were more likely to signal their aggressiveness with either gill displays or attacks rather than retreating immediately. A winning experience then increased the likelihood that the most aggressive behavioral pattern the subordinates exhibited would be attacks rather than gill displays, irrespective of their opponents’ familiarity. Dominants that received a losing experience and faced an unfamiliar opponent were less likely than others to launch attacks directly. And subordinates that challenged dominants with more aggressive tactics but still lost received more post-retreat attacks from their dominant opponents.

Conclusions

Subordinates’ contest decisions were influenced by both their contest experience and the familiarity of their opponents, but these influences appeared at different stages of a contest and did not interact significantly with each other. The influence of a losing experience on dominants’ contest decisions, however, did depend on their subordinate opponents’ familiarity. Subordinates and dominants thus appeared to integrate information from the familiarity of their opponents and the outcome of previous contests differently, which warrants further investigation. The higher costs that dominants imposed on subordinates that behaved more aggressively toward them may have been to deter them from either fighting back or challenging them in the future.

     

Self-organizing dominance hierarchies in a wild primate population

Linear dominance hierarchies, which are common in social animals, can profoundly influence access to limited resources, reproductive opportunities and health. In spite of their importance, the mechanisms that govern the dynamics of such hierarchies remain unclear. Two hypotheses explain how linear hierarchies might emerge and change over time. The ‘prior attributes hypothesis’ posits that individual differences in fighting ability directly determine dominance ranks. By contrast, the ‘social dynamics hypothesis’ posits that dominance ranks emerge from social self-organization dynamics such as winner and loser effects. While the prior attributes hypothesis is well supported in the literature, current support for the social dynamics hypothesis is limited to experimental studies that artificially eliminate or minimize individual differences in fighting abilities. Here, we present the first evidence supporting the social dynamics hypothesis in a wild population. Specifically, we test for winner and loser effects on male hierarchy dynamics in wild baboons, using a novel statistical approach based on the Elo rating method for cardinal rank assignment, which enables the detection of winner and loser effects in uncontrolled group settings. Our results demonstrate (i) the presence of winner and loser effects, and (ii) that individual susceptibility to such effects may have a genetic basis. Taken together, our results show that both social self-organization dynamics and prior attributes can combine to influence hierarchy dynamics even when agonistic interactions are strongly influenced by differences in individual attributes. We hypothesize that, despite variation in individual attributes, winner and loser effects exist (i) because these effects could be particularly beneficial when fighting abilities in other group members change over time, and (ii) because the coevolution of prior attributes and winner and loser effects maintains a balance of both effects.     

Male attractiveness covaries with fighting ability but not with prior fight outcome in house crickets

In many species males that tend to win fights against othermales are more attractive to females. There are three ways inwhich male fighting ability and attractiveness may be associated:(1) attractiveness and fighting ability are influenced by thesame underlying traits (e.g., body size), (2) females prefermales that have directly observed winning fights, or (3) winningprevious fights indirectly improves a male's chance of beingpreferred by females. The last possibility may arise as a consequenceof the "loser effect"; in many species when a male loses a fighthis probability of losing subsequent fights increases. Thereare, however, no studies testing whether such a "loser effect"also influences male attractiveness. Here we show that maleattractiveness and fighting ability are positively correlatedin the house cricket, Acheta domesticus. Our experiment wasdesigned so that females could not directly observe the outcomeof fights, thus eliminating possibility (2) above. We then testedbetween possibilities (1) and (3) by making use of the factthat in some cricket species the "loser effect" can be eliminatedexperimentally by ‘shaking’ a male and stimulatingthe motor program for flying. We showed that in A. domesticus‘shaking’ does affect the outcome of subsequentfights. Males that had won two previous fights were less likelyto win a fight after being ‘shaken’ than when subjectto a control treatment. In contrast, males that had lost twoprevious fights were more likely to win a fight after being‘shaken’ than when they were not shaken. There was,however, no effect of ‘shaking’ on male attractiveness.We conclude that the "loser effect" does not alter the tendencyfor large, dominant males to be attractive to females. Instead,it appears that there are traits correlated with both fightingability and attractiveness. One such trait is body size. Fightwinners were significantly larger than losers and attractivenesswas positively correlated with male body size.     

Previous experience and contest outcome: winner effects persist in absence of evident loser effects in a parasitoid wasp

Abstract The experience of a previous conflict can affect animals' performance during a later contest: a victory usually increases and a defeat usually decreases the probability of winning a subsequent conflict. These winner and loser effects could result from a reassessment by contestants of their perceived fighting abilities. Game-theoretic models based on this assumption predict that a loser effect can exist alone or in the presence of a winner effect, but a winner effect cannot persist alone, at least when contestants are young and without experience of contest. Moreover, when both effects coexist, the loser effect is expected to be of a greater magnitude and last longer than the winner effect. To date, these predictions have been supported by empirical evidence. Here we show for the first time that a winner effect can exist in the absence of any evident loser effect in a parasitoid wasp, Eupelmus vuilleti, when fighting for hosts. This finding consequently raises questions about the possible mechanisms involved and challenges the main assumption of previous theoretical models. We suggest an alternative explanation for the evolution of only winner effects that is based on the modification of contestants' subjective value of the resource rather than on a reestimation of their fighting abilities.     

Traits of winner and loser species indicate drivers of herb layer changes over two decades in forests of NW Germany

Questions: What are the most likely environmental drivers for compositional herb layer changes as indicated by trait differences between winner and loser species? Location: Weser‐Elbe region (NW Germany). Methods: We resurveyed the herb layer communities of ancient forest patches on base‐rich sites of 175 semi‐permanent plots. Species traits were tested for their ability to discriminate between winner and loser species using logistic regression analyses and deviance partitioning. Results: Of 115 species tested, 31 were identified as winner species and 30 as loser species. Winner species had higher seed longevity, flowered later in the season and more often had an oceanic distribution compared to loser species. Loser species tended to have a higher specific leaf area, were more susceptible to deer browsing and had a performance optimum at higher soil pH compared to winner species. The loser species also represented several ancient forest and threatened species. Deviance partitioning indicated that local drivers (i.e. disturbance due to forest management) were primarily responsible for the species shifts, while regional drivers (i.e. browsing pressure and acidification from atmospheric deposition) and global drivers (i.e. climate warming) had moderate effects. There was no evidence that canopy closure, drainage or eutrophication contributed to herb layer changes. Conclusions: The relative importance of the different drivers as indicated by the winner and loser species differs from that found in previous long‐term studies. Relating species traits to species performance is a valuable tool that provides insight into the environmental drivers that are most likely responsible for herb layer changes.     

Male contest investment changes with male body size but not female quality in the spider Nephila clavipes

Animals use rules to adjust their level of investment in a contest. We evaluate male strategies during contests over females in the golden orb-web spider Nephila clavipes. We tested whether male behaviour changes with female value, and found that contests were similar in intensity and outcome whether the female was a juvenile or adult, virgin or non-virgin, or whether one male had invested sperm in the female. We found evidence that males use a self-assessment strategy when deciding to withdraw from a contest. Loser body size and contestant size difference were correlated with a higher frequency of contest escalation, and fights involving two large males were more likely to escalate than a fight in which one male was small. A multiple regression showed that loser body size had a stronger effect on contest escalation than contestant body size difference. More importantly, the size of the winning male had no effect on contest escalation, a key prediction of a self-assessment strategy. In N. clavipes, body size is the primary factor that determines the outcome of male contests, and males do not appear to assess their opponent or the quality of the resource when deciding to withdraw from the fight.     

Juvenile social experience affects pairing success at adulthood: congruence with the loser effect?

Social interactions with adults are often critical for the development of mating behaviours. However, the potential role of other primary social partners such as juvenile counterparts is rarely considered. Most interestingly, it is not known whether interactions with juvenile females improve males’ courtship and whether, similar to the winner and loser effects in a fighting context—outcome of these interactions shapes males’ behaviour in future encounters. We investigated the combined effects of male quality and juvenile social experience on pairing success at adulthood in zebra finches (Taeniopygia guttata). We manipulated brood size to alter male quality and then placed males in either same- or mixed-sex juvenile dyads until adulthood. We found that males from reduced broods obtained more copulations and males from mixed-sex dyads had more complete courtships. Furthermore, independent of their quality, males that failed to pair with juvenile females, but not juvenile males, had a lower pairing success at adulthood. Our study shows that negative social experience with peers during adolescence may be a potent determinant of pairing success that can override the effects of early environmental conditions on male attractiveness and thereby supports the occurrence of an analogous process to the loser effect in a mating context.     

Do female leaf beetles Galerucella nymphaeae choose their mates and does it matter?

The role of active female choice in sexual selection is frequently difficult to ascertain, and this is particularly the case for many insect species. Also, it is uncertain whether choosing between males would affect offspring viability. We designed an experiment to investigate the presence of female choice in a Coleoptera species (Galerucella nymphaeae). We also estimated whether mate choice would have any effect on offspring performance. Females were first placed with two males in a test arena to see which of the males copulated with the virgin female, and how quickly. Subsequently the loser male was offered a new virgin female to test for any change in latency time until mating. The two-male tests indicated that males with wider upper prothoraxes were more likely to mate with the female, and the latency time until mating was shorter when the winner male had relatively long wings. When the loser males were placed singly with females the latency time was not correlated with male size, and was the same as when two males were used. These results suggest that male-male competition is the most likely cause of sexual selection on size, and if females have any preferences they are not very strong. The seemingly passive female strategy may be sufficient to ensure that females mate with the most vigorous males, since in the field several males usually compete for access to each female. Finally, the benefits of female choosiness were estimated to be low and non-significant. The eggs of the winner males were no more likely to hatch, offspring survival into adulthood was no greater among descendants of winner males, and the offspring did not differ in adult size. Received: 3 March 1997 / Accepted: 13 October 1997     

What sets the odds of winning and losing?

Social experience influences the outcome of conflicts such that winners are more likely to win again and losers will more likely lose again, even against different opponents. Although winner and loser effects prevail throughout the animal kingdom and crucially influence social structures, the ultimate and proximate causes for their existence remain unknown. We propose here that two hypotheses are particularly important among the potential adaptive explanations: the 'social-cue hypothesis', which assumes that victory and defeat leave traces that affect the decisions of subsequent opponents; and the 'self-assessment hypothesis', which assumes that winners and losers gain information about their own relative fighting ability in the population. We discuss potential methodologies for experimental tests of the adaptive nature of winner and loser effects.     

On the evolution of pure winner and loser effects: A game-theoretic model

The persistence of linear dominance hierarchies is often attributed to higher probabilities of a win after a win or a loss after a loss in agonistic interactions, yet there has been no theory on the evolution of such prior-experience effects. Here an analytic model, based on the idea that contests are determined by subjective perceptions of resource-holding potential (RHP) which animals may revise in the light of experience, demonstrates that winner and loser effects can evolve through round-robin competition among triads of animals drawn randomly from their population, and that the probability of a hierarchy increases with the strength of the combined effect. The effects are pure, in the sense that a contestant observes neither its own RHP nor its opponent’s RHP or RHP perception or win—loss record; and so the strength of an effect is unmodified by the RHPs of particular individuals, but depends on the distribution of RHP among the population at large. The greater the difference between an individual’s and its opponent’s RHP perception, the more likely it is to win a contest; however, if it overestimates its RHP, then the cost of fighting increases with the overestimate. A winner or loser effect exists only if the fitness gain of the beta individual in a hierarchy, relative to that of the alpha, is less than 0.5. Then a loser effect can exist alone, or it can coexist with a winner effect; however, there cannot exist a winner effect without a loser effect.     

Size,sex and social experience: Experimental tests of multiple factors mediating contest behaviour in a rockpool fish

Aggressive contests amongst conspecifics are important to understand from an ecological and evolutionary perspective as contest dynamics can directly influence individual fitness. For some species, individual attributes such as relative body size closely predict the outcome, intensity and duration of contests, whereas for others, prior social experiences play a key role. However, disentangling the relative effects of individual attributes and social experiences is challenging from an experimental perspective, and because of this, few studies have investigated relative effects of multiple factors. Rockpool fishes have been well studied in terms of factors governing abundance, distribution and community structure, but much less so in terms of contest behaviour. This is surprising because contest dynamics are likely to directly affect the distribution of fishes along the rocky shore, and hence indirectly govern population and community composition. Here, we explored multiple factors potentially influencing contest behaviour in a numerically dominant, resident intertidal fish species, Bathygobius cocosensis (Gobiidae). Using a series of staged pairwise contest trials, we investigated the effect of size, sex and social experiences (namely prior residency and winner–loser experiences) on contest dynamics. We found no evidence that prior residency influenced contest outcome, suggesting social experiences play a minor role in contest dynamics. Previous winner/loser experience also did not influence contest outcome, although this is likely a product of low sample size. In addition, the likelihood of winning was unrelated to contestant sex, and the combination of sexes in paired contests did not influence contest intensity or duration. Instead, body size was related to contest outcome, intensity and duration in the majority of experimental trials. These results suggest that body size rather than sex or social experiences is the key determinant of contest dynamics in this species. We suggest that the dynamic biotic and abiotic environment inhabited by intertidal fish may reduce the influence of prior social experiences in modulating contest dynamics.