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This study examined the relative importance of contest experience and size differences to behavioral decisions over the course of contests. Using a mangrove rivulus fish, Kryptolebias marmoratus, we showed that although contest experience and size differences jointly determined contest outcomes, they affected contestants' interactions at different stages of contests. Contest experience affected behavioral decisions at earlier stages of contests, including the tendency and latency to launch attacks, the tendency to escalate contests into mutual attacks and the outcome of non-escalated contests. Once contests were escalated into mutual attacks, the degree of size difference affected the fish's persistence in escalation and chance of winning, but contest experience did not. These results support the hypothesis that contest experience modifies individuals' estimation of their fighting ability rather than their actual strength. Furthermore, (1) in contests between two naïve contestants, more than 60 % of fish that were 2-3 mm smaller than their opponent escalated the contest to physical fights, even though their larger opponents eventually won 92 % of escalated fights and (2) fish with a losing experience were very likely to retreat in the face of an opponent 2-3 mm smaller than them without escalating. The result that a 2-3 mm size advantage could not offset the influence of a losing experience on the tendency to escalate suggests that, as well as depending on body size, the fish's physical strength is influenced by other factors which require further investigation.

Resource ownership often increases an individual's aggressiveness and its probability of defeating a competitor. Individuals contesting resource owners could therefore incur higher costs, making individuals reluctant to compete with owners. We tested the hypothesis that animals use asymmetry in resource ownership as a cue for contest costs and adjust contest decisions accordingly. Using a mangrove rivulus fish (Kryptolebias marmoratus), we staged (1) contests with a randomly assigned asymmetry in resource ownership (one fish was provided with a shelter) and (2) contests in which neither fish had a shelter. Owners that were in their shelters at the contest start showed a greater tendency to fight and won more contests than their intruder opponents; those outside the shelter at the start did not. Compared with fish in contests with no shelters at stake, shelter owners had a higher tendency to fight whether or not they were in their shelters at the start; intruders, however, had a lower tendency to fight only against owners that were inside the shelter at the start. These results demonstrate (1) that ownership status influences both owners’ and intruders’ contest decisions (and in opposite directions), producing a detectable ownership advantage and (2) that intruders required confirmation of their opponents’ ownership status before retreating without challenging them. Ownership status per se is therefore important to the fish's contest decisions.

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

Animals’ contest performance is influenced by their recent contest experiences. This influence could either be exerted by individuals re-estimating their own fighting ability (self-assessment) or by their opponents responding to status-related cues (social-cue mechanism) or both. Individuals of Kryptolebias marmoratus, a hermaphroditic killifish, were given different contest experiences to examine how two opponents’ prior experiences combined to determine their contest interaction and to test both of these mechanisms as potential causes of the observed experience effect. Our data showed that losers’ decisions to retreat at different stages of a contest were influenced by their own but not by the winners’ contest experience—a result consistent with self-assessment but not with the social-cue mechanism. An association between the fish initiating and winning contests thus probably arose because both were correlated with an individual’s assessment of its fighting ability, but not because initiating contests made opponents more inclined to retreat.

To determine whether animals assess each other's fighting ability in contests, researchers usually regress contest duration over the sizes of the contestants. The predominant trend in recent studies is for the contest duration to correlate positively with the size of the smaller opponent but to have no obvious relationship with the size of the larger opponent. This indicates that animals make contest decisions based on their own abilities (‘self assessment’) and displaces the once-popular belief that they assess their opponents (‘mutual assessment’). These tests, however, are based on the implicit but never stated assumption that animals adopt only one assessment approach throughout an entire contest. By examining the contest behaviours of a killifish, we show that (1) the fish adopt mutual assessment at earlier stages when deciding whether to escalate the contest from the mutual display to the attack phase and (2) once a contest is escalated, the fish switch to self assessment to decide how long to escalate. Our results show that individuals may adopt multiple assessment approaches in one contest; contest behaviours in different stages (where applicable) of a contest should be analysed separately to better elucidate animal contest assessment strategies.

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.

We examined the effect of prior winning and losing experiences on the initiating and responding strategies of contestants in contests between individuals of Rivulus marmoratus (Cyprinodontidae). Each contestant was given a penultimate and a recent fighting experience approximately 48 and 24 h prior to the dyadic contests, respectively, through randomly selected procedures. Winning and losing experience appeared to influence different types of fighting behaviours. Losing experiences decreased the probability of an individual initiating a confrontation and thus increased its tendency to retreat immediately when challenged. Winning experiences did not affect the probability of initiation, but significantly increased the likelihood of an individual initiating with attacks that effectively deterred its opponents. A substantial proportion (59/153) of individuals retreated immediately when challenged and reduced the number of fights available for examining experience effects on responding strategies at later stages of a contest. None the less, winning experiences consistently increased the likelihood of an individual retaliating by attacking its opponent at various stages of a contest, and eventually increased its probability of escalating a confrontation into physical fights. However, the effects of losing experiences on these responding strategies were undetectable. Recent experiences significantly affected all fighting behaviours examined, but penultimate experiences significantly affected only the tendency to initiate a confrontation with attacks and the likelihood of escalation. These results indicated that prior experiences had the longest lasting effect on the potentially most costly fighting behaviour. Prior experiences influenced the outcome of nonescalated contests as well as the probability of escalation, but did not significantly affect the outcome of escalated contests. These results are consistent with the hypothesis that prior experiences modify the information that an individual has about its fighting ability but do not alter its actual fighting ability and that actual fighting ability becomes the more important influence on outcomes of escalated contests.

Aggression plays an important role in animal contests, but the extent to which aggression correlates with dominance has been a topic of much debate. The relationship between aggression and dominance ability in the hermaphroditic fish, Rivulus marmoratus, was investigated using three standard protocols, the mirror test (Mi), model test (Mo), and standard opponent test (So). In each, display latency, attack latency, and biting frequency were quantified for a test individual towards its opponent. The general rank-order for eliciting strength of the three different stimuli was Mi > So > Mo. The relationships between the individual indices from the standard tests and three dyadic contest variables, initiator of display, initiator of attack, and winner, were analysed in contests between previously tested pairs to ascertain how well the standard protocols predicted dyadic contest behaviour/outcome. Display and attack latencies in the standard tests did not predict the level of analogous combat behaviour. Biting frequency differences between individuals in a pair in the So and Mo tests as well as display latency differences in the Mi test contributed to predictions of contest outcome. The individual that scored higher, relative to its opponent, won a significantly greater proportion of the bouts. These findings demonstrate the importance of relative differences in aggression in determining dominance. However, the predictive value of standard test behaviour is test-specific and, based on the available literature, depends on both the species used and the context in which they are employed.

An important question in state-dependent behaviour is how multiple influences on state are integrated to determine current behaviour. Aggressive behaviour is known to be affected by a prior contest experience. Nevertheless, whether and how multiple prior fighting experiences are integrated into a fighting decision remain unexplored. In this study, individuals of Rivulus marmoratus (Cyprinodontidae), a hermaphroditic fish, were given different combinations of two prior fighting experiences to investigate: (1) the effect of penultimate experiences on the probability of winning a subsequent contest; (2) the relative effect of a recent win and loss; and (3) whether the effect of a winning experience was as short lived as observed in other species. Penultimate and recent fighting experiences were given to the test fish approximately 48 and 24 h prior to the dyadic contests, respectively. From the results of the five types of contests staged, we conclude that: (1) penultimate fighting experiences had a significant effect on the probability of winning a subsequent contest; (2) a more recent experience had a more pronounced effect than an earlier experience, which suggested that the effect of a fighting experience would decay and/or the effect of a recent experience would interfere with the effect of an earlier experience; (3) no asymmetric effect between a winning experience and a losing experience was detected; and (4) the effect of both a winning and a losing experience lasted for at least 48 h in R. marmoratus which was the maximum time tested in these experiments. The possible reasons for the differences in results among studies of experience effects on contest outcomes are discussed. Copyright 1999 The Association for the Study of Animal Behaviour.

Many animals modify behavioural decisions based on information they have previously acquired. Contest behaviour is often affected by previous contest experiences: individuals behave more and less aggressively after a victory and defeat, respectively (winner/loser effect). Individuals in the field sometimes encounter multiple competitors in quick succession, but whether these experiences interact to influence each other’s importance is unclear. We tested five hypotheses for experience interaction (no interaction, retroactive interference, proactive interaction, reinforcement and diminishing returns) using Kryptolebias marmoratus. Focal individuals were paired up with opponents having the same 1-month contest outcome (1 month before the experiment), as this difference in actual or perceived fighting ability has been shown to affect the fish’s response to new experiences. We gave the focal individual of a pair a winning or losing experience on day 1. Then both fish of the pair received the same winning, losing or no-contest experience on day 2. Then we organised fights between the two. The effect of a day-1 losing experience did depend on the fish’s actual or perceived fighting ability: one-month losers readily showed loser effects from the day-1 losing experience, irrespective of the day-2 experience (i.e. no interaction between day-1 and day-2 experiences). One-month winners, however, only showed loser effects from a day-1 losing experience when the day-2 experience was also a loss (i.e. reinforcement). Day-1 winning experiences did not interact with day-2 experiences in 1-month losers or winners. Therefore, multiple experiences sometimes reinforce each other, but how they combine to influence behaviour depends on an individual’s actual or perceived fighting ability.

Introduction Animals are capable of using information from recent experiences to modify subsequent behavioral responses. Animals' ability or propensity to modify their behavior in the light of new information has repeatedly been shown to correlate with, or be influenced by, either their intrinsic competitive ability or their dominance experience - an influence which can be long-lasting. Using a mangrove killifish, Kryptolebias marmoratus, as the study organism, we investigated whether and if so how the effect of a winning or a losing experience one day prior to a dyadic contest was modulated by both competitive ability measured two months previously and a winning or losing experience forced on the contestants one month previously. Results Winning/losing experience forced on the fish one month previously affected how they utilized information from their winning/losing experience one day before Test Day: Individuals that were randomly assigned a losing experience one month previously were more susceptible to the influence of their 1-day winning/losing experience than those assigned a winning experience. Competitive ability measured two months previously, winning/losing experience from one month previously and the winning/losing experience received one day previously all significantly influenced the fish's contest behaviors on Test Day, although only 2-month competitive ability significantly influenced escalation duration, indicating that it was still a good index for the fish's competitive ability two months later. Conclusions These results suggest that the value to the fish of information from a recent win or loss depends on the outcome of their past contests and show that contest experience has a long-term effect on contest behavior.

Introduction Many animals use information acquired from recent experiences to modify their responses to new situations. Animals’ decisions in contests also depend on their previous experience: after recent victories individuals tend to behave more aggressively and after defeats more submissively. Although these winner and/or loser effects have been reported for animals of different taxa, they have only recently been shown to be flexible traits, which can be influenced by extrinsic factors. In a mangrove killifish (Kryptolebias marmoratus), for instance, individuals which lost an earlier contest were more likely than others to alter contest decisions after a recent win/loss. This result suggests that individuals perceiving themselves to have worse fighting abilities are more inclined to adjust contest strategy based on new information. If this is the case, an individual’s propensity to modify behaviour after a win/loss might also be modulated by intrinsic mechanisms related to its ability to fight. Stress and sex steroid hormones are often associated with an individual’s contest behaviour and performance, so, in this study, we tested the hypothesis that an individual’s propensity to change behaviour after wins or losses also depends on its hormonal state. Results Our results show that an individual’s propensity to adjust contest decisions after wins and losses does depend on its hormonal state: individuals with lower levels of cortisol (F), testosterone (T) and 11-ketotestosterone (KT) are more receptive than others to the influence of recent contest experiences, especially losing experiences, and the influences last longer. Furthermore, although winning and losing experiences resulted in significant changes in behaviour, they did not bring about a significant change in the levels of F, T, KT or oestradiol (E2). Conclusions This study shows that an individual’s receptivity to the influence of recent wins and losses is modulated by its internal state, as well as by extrinsic factors. Individuals with hormonal profiles corresponding to lower aggressiveness and a reduced likelihood of winning were more likely to alter contest decisions after a recent win/loss. The results also suggest that F, T, KT and E2 are not the primary physiological mechanisms mediating winner-loser effects in this fish.