Assessment Strategies and the Effects of Fighting Experience on Future Contest Performance in the Green Anole (Anolis carolinensis)

Social experiences can be useful sources of information for animals charged with making fitness‐related decisions. Fighting experience can alter an animal's perception of its fighting ability possibly leading to changes in future contest decisions, which may increase/decrease their probability of winning future contests. Winner and loser effects have been revealed in a wide array of animals, but studies using reptilian models are rare. This study investigated the impact of fighting experience on future contest performance and outcome in the green anole lizard and investigated the assessment strategies used by anoles during contests of different intensities. To determine whether the green anole expresses winner or loser effects, focal animals engaged in a primary contest with a smaller (larger) opponent to gain a winning (losing) experience; opponent size asymmetries were a significant predictor of contest outcome. Focal individuals were isolated for 2 d before being given a secondary contest with a size‐matched, naïve opponent. We found no evidence of winner or loser effects 2 d following a previous contest. Although previous contest outcome did not dictate future contest success, dynamics of the previous contest did. Highly aggressive primary contest losers won a significant proportion of the secondary contests, while less aggressive losers were more apt to lose the secondary contest. Secondary contest success of prior winners was not influenced by earlier contest performance. Further analyses of contest dynamics reveal that individuals may use different assessment strategies depending on the intensity of the contest. Our results demonstrate that future contest success may be driven more by individual performance in a prior contest and less by prior contest outcome.     

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.     

Rats Benefit from Winner and Loser Effects

Prior fighting experience of opponents can influence the outcome of conflicts. After a victory, animals are more likely to win subsequent contests, whereas after a defeat animals are more likely to lose, regardless of the identity of opponents. The underlying mechanisms and the adaptive significance of these winner and loser effects are as yet unknown. Here, we tested experimentally whether agonistic behavior of male wild‐type Norway rats is influenced by social experience, and we investigated whether this might reduce fighting costs (duration of contest, risk of injury) in subsequent encounters. Rats were randomly assigned to receive either a losing or a winning experience and subsequently tested with unfamiliar, naïve opponents. We found that most rats with a winning experience won the subsequent encounter, and all rats with a losing experience lost the next contest. Previous winners attacked more rapidly in the subsequent encounter and reduced their aggressive behavior sooner; the contests were decided more quickly, which saved time and behavioral effort to the winner. Previous losers received less aggression in the next encounter, despite emitting fewer submissive ultrasonic calls than in the preceding contest, thereby reducing the risk of being injured by the opponent. Thus, anonymous social experience influenced rats’ subsequent behavior toward size‐matched, naïve, unknown social partners. Furthermore, apparently, they benefit from showing winner and loser effects in intraspecific contests by saving time, energy, and risk.     

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.     

Body Size, Leg Autotomy, and Prior Experience as Factors in the Fighting Success of Male Crab Spiders, Misumenoides formosipes

Game theory predicts which characteristics of male crab spiders should affect their success in mate-guarding contests. Previous data on Misumenoides formosipes suggested that body size and leg number influenced fight outcome, but the study had methodological problems. Using improved methods, we corroborated the size advantage but refuted the disadvantage of leg reduction. We also conducted the first test for an invertebrate on the effect that a single contest has on the outcome of subsequent contests. When size and residency status were controlled, prior winners defeated prior losers after only one contest experience. We discuss the importance of controlling for previous experiences in studies of contest resolution and the ramifications for both past and future studies.     

Modulation of aggressive behaviour by fighting experience: mechanisms and contest outcomes

Experience in aggressive contests often affects behaviour during, and the outcome of, later contests. This review discusses evidence for, variations in, and consequences of such effects. Generally, prior winning experiences increase, and prior losing experiences decrease, the probability of winning in later contests, reflecting modifications of expected fighting ability. We examine differences in the methodologies used to study experience effects, and the relative importance and persistence of winning and losing experiences within and across taxa. We review the voluminous, but somewhat disconnected, literature on the neuroendocrine mechanisms that mediate experience effects. Most studies focus on only one of a number of possible mechanisms without providing a comprehensive view of how these mechanisms are integrated into overt behaviour. More carefully controlled work on the mechanisms underlying experience effects is needed before firm conclusions can be drawn.Behavioural changes during contests that relate to prior experience fall into two general categories. Losing experiences decrease willingness to engage in a contest while winning experiences increase willingness to escalate a contest. As expected from the sequential assessment model of contest behaviour, experiences become less important to outcomes of contests that escalate to physical fighting.A limited number of studies indicate that integration of multiple experiences can influence current contest behaviour. Details of multiple experience integration for any species are virtually unknown. We propose a simple additive model for this integration of multiple experiences into an individual's expected fighting ability. The model accounts for different magnitudes of experience effects and the possible decline in experience effects over time.Predicting contest outcomes based on prior experiences requires an algorithm that translates experience differences into contest outcomes. We propose two general types of model, one based solely on individual differences in integrated multiple experiences and the other based on the probability contests reach the escalated phase. The difference models include four algorithms reflecting possible decision rules that convert the perceived fighting abilities of two rivals into their probabilities of winning. The second type of algorithm focuses on how experience influences the probability that a subsequent contest will escalate and the fact that escalated contests may not be influenced by prior experience. Neither type of algorithm has been systematically investigated.Finally, we review models for the formation of dominance hierarchies that assume that prior experience influences contest outcome. Numerous models have reached varied conclusions depending on which factors examined in this review are included. We know relatively little about the importance of and variation in experience effects in nature and how they influence the dynamics of aggressive interactions in social groups and random assemblages of individuals. Researchers should be very active in this area in the next decade. The role of experience must be integrated with other influences on contest outcome, such as prior residency, to arrive at a more complete picture of variations in contest outcomes. We expect that this integrated view will be important in understanding other types of interactions between individuals, such as mating and predator-prey interactions, that also are affected significantly by prior experiences.     

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.     

Reciprocity between endocrine state and contest behavior in the killifish, Kryptolebias marmoratus

Given the dramatic behavioral effects of winning and losing contests, and pronounced changes in stress and sex steroid hormones post-fight, it is reasonable to suppose that these hormones also dictate future behavior. We sampled water-borne cortisol, testosterone (T), and 11-ketotestosterone (KT) before and after contests in the mangrove killifish, Kryptolebias marmoratus, to determine how endogenous steroid hormone levels might predict and respond to contest dynamics or success. Pre-fight cortisol related negatively, and pre-fight T related positively to contest initiation and winning, particularly in the smaller opponent. In the pairs where a larger fish won the contest, winners with higher pre-fight T and lower pre-fight cortisol delivered more attacks to the losers. Contest duration and escalation influenced post-fight hormone concentrations primarily in losers. Escalation significantly increased post-fight cortisol, T, and KT for losers but not for winners. However, winners that attacked losers at higher rates had higher levels of post-fight cortisol. Losers also demonstrate the most consistent post-fight hormone responses, particularly to contest escalation and duration. Despite the bidirectional relationship between hormones and contest behavior, we found no overall mean differences in pre- or post-fight cortisol, T, or KT between eventual winners and losers. Thus, it is evident that the categorical states of winner and loser cannot alone reveal the complex, reciprocal associations between endocrine systems and social behavior.     

Effect of body weight, antler length, resource value and experience on fight duration and intensity in fallow deer

We tested predictions of evolutionary game theory focusing on fight duration and intensity during contests between European fallow deer, Dama dama L. We examined the relation between contest duration and intensity and resource-holding potential (RHP; body weight and antler size), in an effort to reveal the assessment rules used by competing males. We examined other potential determinants of duration and intensity: resource value (the oestrous female) and experience of agonistic interactions. Asymmetry in body weight or antler length of contestants was not correlated with fight duration. Body weight and antler length of the fight winner or loser were also not correlated with fight duration. Neither were the body weight of the heavier or lighter animal or the antler length of the animal that had longer or shorter antlers. A measure of intensity (the jump clash) was positively related to the body weight of the losing animal and the lighter member of the dyad. These results are consistent with the hypothesis that opponents escalate contest intensity based on assessment of their own ability rather than through mutual assessment. There was no evidence that resource value is an important factor in either fight duration or intensity in this population. As the number of fights between pairs of males increased, there was a decrease in fight duration. Fights were longer when at least one member of a competing pair of males had previously experienced a victory.     

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.     

Influence of previous residency and body mass in the territorial contests of the butterfly Hermeuptychia fallax (Lepidoptera: Satyrinae)

Although many studies have evaluated the possible rules used by males to settle territorial contests, few have simultaneously considered the possibility of mutual or self-assessment of fighting ability and the effect of prior residence on the chances of winning. Here, we experimentally investigated the influence of previous residency on the development and outcome of contests between males of the butterfly Hermeuptychia fallax. By comparing the frequency of resident victories in nonmanipulated disputes versus interactions in which both rivals were induced to behave as residents, we showed that prior residency may interact with body mass in contest resolution. In nonmanipulated disputes, resident males showed high winning probabilities, although they were similar to losers in body mass. However, when both males behaved as residents, the disputes were longer and were settled in favor of the heavier male. This indicates that, although residency is an important component of male willingness to fight, it is not the only factor influencing conflict resolution. Despite the correlation between body mass and territorial success, the weight of neither winners nor losers correlated with fighting duration. We conclude that residence asymmetry may sometimes conceal male traits important to contest resolution and that it may obscure the identification of the assessment strategy used in contests.     

Victory displays: a game-theoretic analysis

Two rationales have been proposed verbally for the functionof victory displays, which are performed by the winners of contestsbut not by the losers. The "advertising" rationale is that victorydisplays are attempts to communicate victory to other membersof a social group that do not pay attention to contests or cannototherwise identify the winner. The "browbeating" rationale isthat victory displays are attempts to decrease the probabilitythat the loser of a contest will initiate a future contest withthe same individual. We formally explore the logic of theserationales with game-theoretic models. The models show thatboth rationales are logically sound; however, all other thingsbeing equal, the intensity of victory displays will be highestthrough advertising in groups where the reproductive advantageof dominance is low and highest through browbeating in groupswhere the reproductive advantage of dominance is high.     

Ultraviolet signals fighting ability in a lizard

Ultraviolet (UV) signals are used in female mate choice in numerous taxa; however, the role of UV signals in male contests remains relatively unexplored. We experimentally reduced throat UV of free-ranging lizards (Platysaurus broadleyi) to test whether UV acts as a signal of fighting ability during male contests. We found that UV-reduced males were more likely to be challenged than control males. However, contest outcome was not influenced by UV-reduction, and this was despite other obvious asymmetries between opponents, such as body size and residency. Throat UV was confirmed as a signal of fighting ability because contests were more likely to escalate when one contestant had reduced UV. Therefore, throat UV, not body size or residency, was used during the initial stage of opponent assessment, but this did not influence contest outcome. The results suggest that UV overrides other traits that could function as signals during rival assessment.     

The ‘home advantage’ is necessary for a full winner effect and changes in post-encounter testosterone

Winning aggressive contests can both enhance future winning ability and change post-encounter hormones; however, it remains unclear if the context of a fight also influences such winner effects and hormone changes. We investigated this issue by using California mice (Peromyscus californicus) to test if the effect of residency status is necessary to improve future winning ability and alter post-encounter hormones. Male mice were subjected to an aggressive contest and their blood was collected 45 min after the fight. Upon contest initiation, focal mice had a ‘home advantage’ and three prior winning experiences, only one of these factors, or neither factor. Only individuals with a ‘home advantage’ and prior winning experience showed a full winner effect. Post-encounter changes in testosterone and progesterone resulted from an interaction between residency status and winning experience. These data indicate that in male California mice a ‘home advantage’ is necessary to produce the full winner effect and that residency status helps regulate inter-individual variation in the expression of post-encounter testosterone pulses and progesterone changes. Furthermore, these respective behavioral and physiological phenomena might be modulated in a context-specific manner, in part by the surrounding physical environment.     

Effects of body size and resource value on fighting behaviour in a jumping spider

Fights between male Euophrys parvula, a New Zealand salticid spider, consist of a number of discrete ‘stages’, of increasing intensity. Two aspects of these contests were investigated: (1) relative body size of the two opponents, and (2) the presence of a ‘resource’ (a female model). In 92% of all contests where a size difference existed, the larger of the two opponents won. Contest intensity (measured as the highest intensity behaviour elicited during the contest) was inversely correlated with relative body size of the two opponents (measured as carapace width). Contests escalated further when the female model was present. No relationship was found between contest duration and either relative body size or the presence of the model. It is suggested that for contests in which a number of behaviours of different intensity are used, contest intensity may be a better estimate of contest cost than duration. The results are discussed in terms of theoretical models of contest behaviour.     

Determinants of Dominance in the Tree Lizard Urosaurus ornatus: the Relative Importance of Mass,Previous Experience and Coloration

A determination of some of the factors that predict the outcome of contests between male tree lizards, Urosaurus ornatus, was made using logistic regression modelling on matched-pair data. Two-day-long encounters were staged between pairs of males differing in size (snout-vent length and mass), previous contest status (previous winners and previous losers), and coloration (dorsal coloration during their previous contest and throat coloration, a fixed trait). Mass proved to be the best single predictor of contest outcome, resulting in an 80% correct classification rate for predicting winners and losers, far better than the less than 57% correct classification rate for snout-vent length. Previous social status (winner or loser) also was a powerful single predictor of contest outcome with a 793% correct classification rate, as was previous dorsal coloration (76.7%). When combined, mass and previous status produced the strongest combination of predictors with a better than 86% correct classification rate. Contrary to several previous studies, which implicated throat coloration as an important status signal of dominance, our results failed to show that throat coloration is a strong predictor of contest outcome. Possible reasons for this discrepancy with earlier findings are discussed. The logistic regression models also allow prediction of the magnitude of difference in mass between two contestants for there to be an equal chance of winning, given a second asymmetry in contest predictors.     

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.     

Boldness predicts social status in zebrafish (Danio rerio)

This study explored if boldness could be used to predict social status. First, boldness was assessed by monitoring individual zebrafish behaviour in (1) an unfamiliar barren environment with no shelter (open field), (2) the same environment when a roof was introduced as a shelter, and (3) when the roof was removed and an unfamiliar object (Lego® brick) was introduced. Next, after a resting period of minimum one week, social status of the fish was determined in a dyadic contest and dominant/subordinate individuals were determined as the winner/loser of two consecutive contests. Multivariate data analyses showed that males were bolder than females and that the behaviours expressed by the fish during the boldness tests could be used to predict which fish would later become dominant and subordinate in the ensuing dyadic contest. We conclude that bold behaviour is positively correlated to dominance in zebrafish and that boldness is not solely a consequence of social dominance.     

Food,fights, and burrows: The adaptive significance of intraspecific aggression in the bay goby (Pisces: Gobiidae)

Summary The effects of size, sex, bout initiation, past experience, and prior residence were examined with respect to the outcome of intraspecific agonistic contests in the bay goby (Lepidogobius lepidus). In addition, the adaptive significance of this behavior was examined by investigating the relationship between dominance and access to two potentially limiting resources; food and the burrows of invertebrate symbionts.For heterosexual and male homosexual contests, size and initiation significantly affected bout outcome. Gender did not affect heterosexual encounters. In female homosexual bouts initiation played a significant role in determining contest outcome while size did not. Both past experience and prior residency had significant effects on the outcome of intraspecific encounters.Dominant gobies had increased access to simulated invertebrate burrows and limited food. It appears that intraspecific aggression is an adaptive phenomenon through which access to limiting resources may be mediated.Portions of this research were completed during my stay as a visiting scientist at the National Marine Fisheries Service, Southwest Fisheries Center, Tiburon, CA