Men’s Interest in Allying with a Previous Combatant for Future Group Combat

Intra- and intergroup conflict are likely to have been recurrent features of human evolutionary history; however, little research has investigated the factors that affect men’s combat alliance decisions. The current study investigated whether features of previous one-on-one combat with an opponent affect men’s interest in allying with that opponent for future group combat. Fifty-eight undergraduate men recruited from a psychology department subject pool participated in a one-on-one laboratory fight simulation. We manipulated fight outcome (between-subjects), perceived fighter health asymmetry (within-subjects), and the presence of a witness (within-subjects) over six sets of five rounds of fighting. Following each set, we asked men how interested they would be in allying with their opponent for future group combat. We found that men were more interested in allying with their opponent for future group combat if their opponent won the fight or if a witness was present, but perceived fighter-health asymmetry did not affect men’s decision to ally with their opponent. Exploratory analyses revealed a two-way interaction between fight outcome and the presence of a witness, such that winners without a witness present expressed less interest in allying with their opponent for future group combat. Our findings suggest that men attend to the benefits of allying with a man who has demonstrated relatively superior fighting ability. Alliance with a previous opponent for group combat may vary with the relationship value of the opponent and the utility of demonstrating cooperativeness to third-party observers. These findings inform our understanding of coalition formation.     

Assessment strategy and the evolution of fighting behaviour

The view is examined that the adaptive value of conventional aspects of fighting behaviour is for assessment of relative RHP (resource holding power) of the combatants. Outcomes of aggressive disputes should be decided by each individual's fitness budget available for expenditure during a fight (determined by the fitness difference between adoption of alternative strategies, escalation or withdrawal without escalation) and on the rate of expenditure of the fitness budget if escalation occurs (determined by the RHPs of the combatants). Thus response thresholds for alternative strategies (“assessments”) will be determined by natural selection on a basis of which opponent is likely to expend its fitness budget first, should escalation occur. This “loser” should retreat (before escalation) and the winner should stay in possession of the resource. Many aggressive decisions depend on whether one is a resource holder, or an attacker. Assuming the RHP of the combatants to be equal, there are many instances of fitness pay-off imbalances between holder and attacker which should weight the dispute outcome in favour of one or other opponent by allowing it a greater expendable fitness budget. Usually the weighting favours the holder; the attacker therefore needs a correspondingly higher RHP before it may be expected to win. This is not invariably the case, and much observed data fits the predictions of this sort of model. If assessments are perfect and budget expenditure rates exactly predictable, then there would never seem to be any case for escalation. Escalation can be explained in terms of injury inflictions (expenditures) occurring as discrete events; i.e. as “bouts” won or lost during fighting. Assessment can give only a probabilistic prediction of the outcome of a bout. A simple model is developed to investigate escalation situations. Each combatant assesses relative RHP; this correlates with an absolute probability of winning the next bout (c_abs). The stake played for is infliction of loss of RHP and is determined by the fitness budgets of the opponents. (Each individual plays for the withdrawal of its opponent.) This defines a critical probability of winning (c_crit) for each combatant, above which escalation is the favourable strategy (c_abs > c_crit) and below which withdrawal is favourable (c_abs < c_crit). Escalation should occur only where c_abs-c_crit is positive for both combatants. This model gives predictions compatible with the observations, indicating that RHP loss alone can be adequate to explain withdrawal: escalation behaviour. Withdrawal tendency will be increased by low searching costs. Escalations should be restricted to closely matched RHP opponents if RHP disparity is the major imbalance. Outside the “escalation range” of a given individual, the higher RHP individual wins and the lower one loses (i.e. it should withdraw after conventional display). RHP disparity and holder: attacker imbalance should both interact to shape the observed pattern, though their relative importances will depend on species and situation. In some instances selection may favour immediate withdrawal from an occupied territory even without assessment of RHP.     

Is facial structure an honest cue to real-world dominance and fighting ability in men? A pre-registered direct replication of

Masculine facial morphology (e.g., larger jaw, prominent cheekbones) have been linked to a suite of social outcomes—including greater wealth, career progress, romantic desirability, and even greater political success. A leading explanation for these links is that dominant facial structures represent honest cues of physical dominance and fighting ability. T?ebický et al. (2013) published the first study to demonstrate that masculine facial cues (e.g., large nose, deep-set eyes) similarly predict both dominance judgments and real-world fighting success, but no studies have replicated these popular findings that a large and specific assortment of masculine facial structures are implicated in both fighting ability and dominance perception. Thus, we conducted a pre-registered direct replication and extension of T?ebický et al. (2013). Two separate samples of United States MTurk participants rated 516 UFC fighters' facial photographs on perceived aggressiveness (N = 500) and fighting ability (N = 500). Results showed that perceived aggressiveness was associated with masculine facial morphology (e.g., large nose, deep-set eyes) independent of bodily size. There was also some evidence that perceived fighting ability was associated with masculine facial morphology, which disappeared after controlling for bodily size. There was no support for the relation between facial structure and fighting success, and there were often negative relations between perceived aggressiveness and fighting ability on fighting success (i.e., more successful fighters were perceived as less successful and aggressive). We argue that our evolved psychology differentially processes the distinct avenues to victory that constitute overall fighting success (e.g., knockout versus submission wins).     

Detailed analysis of fighting in polecats (Mustelidae) using ciné film

The phenomenon of fighting was investigated under laboratory conditions using polecats ( Mustela putorius, M. furo and interspecific hybrids). Diadic interactions between male polecats were recorded on 16 mm cine film in an unfamiliar area of 16 m², one individual having been introduced ten minutes before its opponent.
The behaviour patterns involved in fighting are described and their frequencies of occurrence, duration and average bout lengths specified. Biting, which occupied 41% of the animal's time in fighting, and attacking, which made up 27% of the bouts of behaviour, were the most important actions involved in fighting. Polecats most commonly bite their opponent's neck and bites in this region were the longest in duration. Success in gaining a bite was influenced by the opponent's behaviour at the time of the attack.
Polecats fight when one individual bites its opponent and the opponent retaliates by biting; bite was shown to be the commonest response to being bitten. The length of a fight appears to be determined by the levels of motivation to bite of the two opponents, while biting and being bitten operate as a positive feed back mechanism. The fight terminates when the more aggressive opponent ceases to make spontaneous attacks. The reasons why one individual wins a fight are that it consistently spends more time than its opponent in biting, it persists for longer in making spontaneous attacks and its method of attacking is more efficient.     

Contest competition and men's facial hair: beards may not provide advantages in combat

In contemporary human societies, where direct male-male competition is reduced compared to ancestral societies, sporting competitions remain an avenue for status acquisition via intra-sexual competition. Beards are the most visually salient and sexually dimorphic of men's secondary sexual traits and play a strong role in communicating masculinity, dominance and aggressiveness intra-sexually. Hypotheses have been advanced that beards provide advantages in intra-sexual combat, as protective organs and honest signals of fighting ability. Here we provide the first test of these hypotheses using data from professional mixed martial arts fighters competing in the Ultimate Fighting Championship. We explored whether secondary sexual traits (height, weight, beardedness), fighting stance (southpaw, orthodox), arm reach and past contest experiences impact on contest outcomes. If beards function as protective organs, bearded fighters should succumb to fewer knock-outs, and hence protection to injuries to the jaw, fewer abrasions and lacerations to the face and brain damage than clean-shaven fighters. Alternatively, if beards signal fighting ability then bearded fighters should win more fights. We found no evidence that beardedness was associated with fewer losses by knock-out or greater fighter ability. While fighters with longer reaches won more fights, neither stance nor past experience influenced fight outcomes. We suggest that beards represent dishonest signals of formidability that may serve to curtail the escalation of intra-sexual conflict through intimidation rather than providing advantages in direct combat.     

The Importance of Physical Strength to Human Males

Fighting ability, although recognized as fundamental to intrasexual competition in many nonhuman species, has received little attention as an explanatory variable in the social sciences. Multiple lines of evidence from archaeology, criminology, anthropology, physiology, and psychology suggest that fighting ability was a crucial aspect of intrasexual competition for ancestral human males, and this has contributed to the evolution of numerous physical and psychological sex differences. Because fighting ability was relevant to many domains of interaction, male psychology should have evolved such that a man’s attitudes and behavioral responses are calibrated according to his formidability. Data are reviewed showing that better fighters feel entitled to better outcomes, set lower thresholds for anger/aggression, have self-favoring political attitudes, and believe more in the utility of warfare. New data are presented showing that among Hollywood actors, those selected for their physical strength (i.e., action stars) are more likely to believe in the utility of warfare.     

If animals know their own fighting ability, the evolutionarily stable level of fighting is reduced

We consider a version of the Hawk-Dove game in which an animal knows its own fighting ability but not the ability of its opponent. For this game at evolutionary stability there is a critical level of ability such that animals with ability greater than the critical level play Hawk and animals with ability below the critical level play Dove. We define the level of fighting to be the probability of a Hawk-Hawk fight when two opponents meet. We show that even if an animal does not know the ability of its opponent, knowing its own ability results in a lower level of fighting at evolutionary stability than is found in the standard Hawk-Dove game in which there are no differences in ability or abilities are not known.     

Estimating the energetic cost of fighting in shore crabs by noninvasive monitoring of heartbeat rate

After establishing shore crabs, Carcinus maenas, individually in separate aquaria, we used a noninvasive infrared phototransducer to monitor their heartbeat rate continously before, during and after fights with intruder crabs. We confirmed that heartbeat rate is a reliable indicator of oxygen consumption and then used it to estimate indirectly the energetic cost of fights differing in duration and intensity, and its dependence on prior residence and relative size of opponent. Prior residence in aquaria significantly increased the probability that crabs would initiate fights against intruders. The majority of fights were resolved by aggressive contacts, display being used extensively only against smaller intruders. Fights between evenly sized opponents and between residents and larger intruders involved almost continuous aggression, whereas fights with smaller intruders involved several shorter bouts of aggression. Fight duration was weakly correlated with the relative size of opponents. Heartbeat rate, measured only in residents, was elevated above resting levels throughout fights, hence energy expenditure during fighting increased linearly with fight duration. Contrary to expectation, heartbeat rate was not significantly influenced by relative size of the opponent or by the intensity of aggression. After fighting, heartbeat rate usually returned to resting levels within 30-60 min, recovery taking longer in fights against larger intruders, when the fight was always lost. We propose that prolonged elevation of heartbeat rate in residents that had lost to larger intruders represented a state of alertness, adaptive against impending risks of resource loss or injury. Copyright 2000 The Association for the Study of Animal Behaviour.     

Chemoreception and the Assessment of Fighting Abilities in the Lizard Liolaemus monticola

When an individual faces the risk of a conflict, its ability to make ‘correct’ decisions is crucial to its fitness. Research on decision making has focused mainly on visual and acoustic signals, while chemical signals have received much less attention, despite their relevance for many species. Chemosignals can be detected in the absence of the signaller and, in the context of fighting risk, this property confers the advantage that the receiver can avoid agonistic interactions or, if they are unavoidable, that it can prepare itself for the conflict. I studied the behaviour of males of the lizard Liolaemus monticola in the laboratory when they were confronted with chemosignals of a potential opponent. During this ‘pre‐confrontation’ stage, I tested the following predictions: (1) lizards can derive precise information from chemosignals of conspecifics, and use this to respond with precision to the perceived risk and (2) the best predictor of the receiver behaviour, and therefore the best predictor of the risk involved in the fight, is the relative fighting ability of opponents. As a measure of fighting ability, I used body size. ‘Intruders’ were placed in the terrarium of unfamiliar ‘residents’ during the absence of the latter, and their behaviours were recorded. Simple regressions were performed between the different behavioural variables and with the body sizes of intruder and resident, and with the relative difference in body sizes of opponents. The latter was the best predictor of intruder behaviour: it was negatively correlated with behaviours associated with activity (i.e. motion time), chemoexploration (i.e. number of tongue flicks) and behaviours associated with social interactions (i.e. head bobs). These results suggest that males can process information from chemosignals and decisions made during the ‘pre‐confrontation’ stage are based on the assessment of the relative fighting abilities (i.e. relative body size) of opponents.     

Does Lateral Presentation of the Palmate Antlers During Fights by Fallow Deer (Dama dama L.) Signify Dominance or Submission?

A central aim of the study of animal communication is to identify the mode and content of information transferred between individuals. The lateral presentation of the antler palm between male fallow deer has been described as either a signal of individual quality or an attempt to avoid fighting. In the first case two phenotypic features have been proposed by which transmission of individual quality may be facilitated. These are antler size and antler symmetry. The alternative hypothesis proposes that the lateral presentation of antlers occurs as a consequence of averting a threatening posture and may signify a reluctance to fight. We examined whether mature fallow deer use lateral palm presentation as a display during fights to indicate antler size and symmetry. We found no relationship between presentation rate of the antler and antler size and symmetry. Furthermore, males did not preferentially present their larger antler to their opponent. We also investigated whether the rate at which males presented antlers laterally during a fight was related to their ability to win the fight. Our results show that the male who performed more presentations during a fight was more likely to lose it. There were behavioural differences in the way in which a bout of presentation ended; subsequent losers tended to turn their body away from their opponent and subsequent winners tended to lower their antlers to an opponent which we interpret as an invitation to continue fighting. We conclude that the lateral palm presentation serves to de-escalate fighting between mature fallow deer. It is not a mechanism by which to communicate individual quality but rather an indication that a male is less committed to continuing investment in the current contest.     

The aggressive behaviour of individual male polecats (Mustela putorius, M. furo and hybrids) towards familiar and unfamiliar opponents

Experiments are described which were designed to investigate the significance of individual differences in aggressiveness between polecats and the relation between the familiarity of the opponent and the pattern and outcome of fighting.
The behaviour of Mustela putorius, M. furo and hybrids between the two species was investigated in an indoor arena of 16 m².
Two kinds of fighting between male polecats were recorded, "companion fighting" between cage mates, in which biting was inhibited and neither of the opponents became intimidated and "uninhibited fighting" between unfamiliar individuals from which a winner and loser generally emerged and a rank order formed.
Individual differences in aggressiveness were assessed by means of a scoring system which statistical analysis showed, generally, to be consistent for any one individual. The rank of an animal, however, was found not to be invariably related to its aggression score.
A number of other factors influencing fighting were investigated, the earlier introduction of one individual into the arena increases its chances of winning a fight; when offered a choice, male polecats fight strangers in preference to cage mates; and the separation of a group of cage mates for as little as 48 hours induces them to behave towards one another like strangers with the result that a rank order is formed. Factors which had no apparent effect on fighting between males were the weight of the individual or the presence of females.
The attributes of familiar and unfamiliar opponents are discussed in the light of these findings.     

Offenders tend to be heavier: experimental encounters in mangrove-dwelling monitor lizards (<Emphasis Type="Italic">Varanus indicus</Emphasis>)

The evolution of male-biased sexual size dimorphism is often explained by sexual selection providing competitive advantage to the larger males. The aggressive interactions are often dangerous and energy consuming; thus, it is advantageous to reduce the risks by adjusting behavior to correspond with body size as a predictor of fighting success. Organization of contests into distinct phases with the initial displays preceding the real combat allows individuals to assess the body size and strength of the rival. We staged interactions between mangrove-dwelling monitor lizards (Varanus indicus) to uncover the initialization of aggression and factors determining the course of an encounter. The analyses revealed the importance of both absolute and relative body size of encountering males. The attack rate increases with the body weight of the lizard and offenders initializing a contact phase of the fight tend to be the heavier male of the dyad. Regardless of the final outcome of the combat, the results show that only short visual contact provides sufficient information about the body size of the opponent. This enables combatants to determine whether to initiate the fight or not. This finding together with the ethological details of contests provides the first evidence for the ability of mutual assessment in varanids.     

Pitch lowering enhances men's perceived aggressive intent,not fighting ability

Voice pitch is the primary perceptual correlate of fundamental frequency (f_o) and describes how low or high a voice is perceived by listeners. Prior research showed that men whose habitual voice pitch is lower are perceived to have stronger fighting ability. However, voice pitch is also flexible and can thus be used facultatively to signal states that change situationally, such as current aggressive intent (i.e., readiness to use aggression). Drawing on motivation-structural-rules theory, this research tests the hypothesis that male speakers will be perceived as more likely to attack when they lower (compared to raise) their pitch to address an adversary in a conflict situation. Three experiments using male speakers and listeners supported this hypothesis, both with and without controlling for the perception of the speakers' fighting ability. In contrast, the same experiments found no evidence that pitch lowering enhanced the speakers' perceived fighting ability after controlling for their perceived aggressive intent. Moreover, we found mixed evidence that the speakers' perceived physical strength interacted with pitch modulation to influence their perceived aggressive intent. On balance, these findings show that, at least for men, pitch modulation is primarily an aggressive-intent signal assessed independently of signalers' fighting ability. Future research should distinguish between perceptions of aggressive-intent and fighting-ability when examining the perceptual effects of male voice-pitch modulation in intrasexual competition.     

Draglines and Assessment of Fighting Ability in Cannibalistic Jumping Spiders

The frequency of injury and death during female-female aggression varies in the jumping spider genus Portia, with interactions being more violent (likely to end in death or injury of one of the combatants) in P. labiata (from Sri Lanka) than in another two species (P. fimbriata from Australia and P. schultzi from Kenya). To investigate the role of draglines in the assessment of fighting ability, two types of tests were carried out: 1) dragline discrimination and 2) mirror image response (Portia's reaction to mirror images is comparable to interaction with conspecific rivals). For both types of testing, triplets of equal-size conspecific females were used: one female (the test spider) was exposed to draglines of two equal-size conspecific females they had not encountered before (donor spiders). The fighting abilities of donor spiders were determined directly by staging intraspecific contests between them. In dragline-discrimination tests (spider placed in petri dish containing draglines from two conspecific females with different fighting ability), females of P. labiata, but not the other two species, avoided draglines of the superior fighter (i.e., they spent the majority of their time on draglines of donor spiders with lesser fighting ability). For mirror-image testing, the test spider was placed in a petri dish containing a mirror and draglines. Each test spider was tested on two successive days, with donor draglines in the two tests coming from conspecific females with different fighting ability. In these tests, females of P. labiata (but not the other two species) spent less time embracing (each spider pressing its forelegs, palps and front of body against the other spider) and more time in a part of the petri dish where view of the mirror was obstructed when on the draglines of donor spiders with greater fighting ability than when on the other conspecific's draglines. Findings from this study suggest that P. labiata females use signpost cues associated with draglines to assess the relative fighting abilities of unknown opponents.     

Is boldness a resource-holding potential trait? Fighting prowess and changes in startle response in the sea anemone,Actinia equina

Contest theory predicts the evolution of a stable mixture of different strategies for fighting. Here, we investigate the possibility that stable between-individual differences in startle-response durations influence fighting ability or 'resource-holding potential' (RHP) in the beadlet sea anemone, Actinia equina. Both winners and losers showed significant repeatability of pre-fight startle-response durations but mean pre-fight startle-response durations were greater for eventual losers than for eventual winners, indicating that RHP varies with boldness. In particular, individuals with short startle responses inflicted more attacks on their opponent. Both repeatability and mean-level responses were changed by the experience of fighting, and these changes varied with outcome. In losers, repeatability was disrupted to a greater extent and the mean startle-response durations were subject to a greater increase than in winners. Thus, following a fight, this behavioural correlate of RHP behaves in a way similar to post-fight changes in physiological status, which can also vary between winners and losers. Understanding the links between aggression and boldness therefore has the potential to enhance our understanding of both the evolution of animal personality and the 'winner and loser effects' of post-fight changes in RHP.     

A high aggression strategy for smaller males

Male-male conflict is common among animals, but questions remain as to when, how and by whom aggression should be initiated. Factors that affect agonistic strategies include residency, the value of the contested resource and the fighting ability of the two contestants. We quantified initiation of aggression in a fish, the desert goby, Chlamydogobius eremius, by exposing nest-holding males to a male intruder. The perceived value of the resource (the nest) was manipulated by exposing half of the residents to sexually receptive females for two days before the trial. Resident male aggression, however, was unaffected by perceived mating opportunities. It was also unaffected by the absolute and relative size of the intruder. Instead resident aggression was negatively related to resident male size. In particular, smaller residents attacked sooner and with greater intensity compared to larger residents. These results suggest that resident desert goby males used set, rather than conditional, strategies for initiating aggression. If intruders are more likely to flee than retaliate, small males may benefit from attacking intruders before these have had an opportunity to assess the resident and/or the resource.     

Eavesdropping on visual cues in green swordtail (Xiphophorus helleri) fights: a case for networking

Aggressive contests probably occur in networking environments where information about fighting ability is conveyed both to an opponent and to individuals peripheral to the fight itself, the bystanders. Our primary aim was to investigate the relative influences of eavesdropping and prior social experience on the dynamics of aggressive contests in Xiphophorus helleri. A bystander's ability to witness an encounter was manipulated using clear, one-way mirror, and opaque partitions. After watching (or not watching) the initial contest, the bystander encountered either the winner or loser of the bout. Treatment comparisons of bystander-winner or bystander-loser contest dynamics indicated the presence or absence of winner, loser, or eavesdropping effects. Winner and loser effects had negligible influences on bystander contest dynamics. Eavesdropping significantly reduced the bystander's propensity to initiate aggression, escalate, and win against seen winners regardless of whether the watched bout had escalated or not. Though eavesdropping had relatively little effect on bystander-loser contest dynamics, bystanders were less prone to initiate aggression and win against losers that had escalated in the witnessed bout. Thus, bystanders appear to preferentially retain and utilize information gained about potentially dangerous opponents (winners or persistent losers). Our data lend clear support for the importance of eavesdropping in visually based aggressive signalling systems.     

Superior fighters make mediocre fathers in the Pacific blue-eye fish

It is widely assumed that male competition and female choice select for elaboration of the same male traits and that fighting ability is synonymous with high quality in terms of benefits to females. Under these assumptions, females are expected to use the same traits that reflect fighting ability to choose the most dominant male, even if females are not privy to actual male-male interactions. Few studies, however, have explicitly investigated female choice in relation to male fighting ability. I conducted experiments separating the effects of male competition and female choice in a freshwater fish, the Pacific blue-eye, Pseudomugil signifer, to test whether females prefer dominant males and whether females obtain higher egg hatching success by being choosy. When females were precluded from witnessing agonistic encounters between two potential mates, they did not appear to use traits correlated with fighting ability to choose competitively superior males. However, even when females were privy to competition, witnessing male interactions did not induce a preference for dominant individuals. Lack of preference for superior fighters may be because there was no difference in hatching success between eggs guarded by dominant and subordinate males. Instead, females appeared to prefer males that spent a greater proportion of time engaged in courtship and, in so choosing, enjoyed higher egg hatching success. These results indicate that dominant males are not necessarily more attractive than subordinates nor do the former necessarily guarantee or deliver the kind of benefits that females may seek.     

Fight tactics in wood ants: individuals in smaller groups fight harder but die faster

When social animals engage in inter-group contests, the outcome is determined by group sizes and individual masses, which together determine group resource-holding potential ('group RHP'). Individuals that perceive themselves as being in a group with high RHP may receive a motivational increase and increase their aggression levels. Alternatively, individuals in lower RHP groups may increase their aggression levels in an attempt to overcome the RHP deficit. We investigate how 'group RHP' influences agonistic tactics in red wood ants Formica rufa. Larger groups had higher total agonistic indices, but per capita agonistic indices were highest in the smallest groups, indicating that individuals in smaller groups fought harder. Agonistic indices were influenced by relative mean mass, focal group size, opponent group size and opponent group agonistic index. Focal group attrition rates decreased as focal group relative agonistic indices increased and there was a strong negative influence of relative mean mass. The highest focal attrition rates were received when opponent groups were numerically large and composed of large individuals. Thus, fight tactics in F. rufa seem to vary with both aspects of group RHP, group size and the individual attributes of group members, indicating that information on these are available to fighting ants.     

Analysis of repeated signals during shell fights in the hermit crab Pagurus bernhardus

Shell exchanges between hermit crabs may occur after a period of shell rapping, when the initiating or attacking crab brings its shell rapidly and repeatedly into contact with the shell of the non-initiator or defender, in a series of bouts. There are two opposing models of hermit crab shell exchange and the function of shell rapping. The negotiation model views shell exchange as a mutualistic activity, in which the initiator supplies information about the quality of its shell via the fundamental frequency of the rapping sound. The aggression model views shell rapping as either detrimental to the defending crab, or as providing it with information about the initiator''s ability or motivation to continue, or both. The negotiation model makes no predictions about the temporal pattern of rapping, but under the aggression model it would be expected that crabs that rapped more vigorously would be more likely to effect an exchange. Repeating the signal could be expected under either model. Crabs that achieve an exchange rap more vigorously, rapping is more persistent when a clear gain in shell quality may be achieved, and the vigour is greater when the relative resource-holding potential (or ''fighting ability'') is high. These findings support the aggression model rather than the negotiation model. Contrary to the predictions of game theory, crabs that do not effect an exchange appear to signal that they are about to give up. The data suggest that rapping is performed repeatedly because the accumulation of all of the performances acts as a signal of stamina.