Female Contests for Nest Sites and Mates in the Pied Flycatcher Ficedula hypoleuca

Predictions of game theory models about the outcome of animal conflicts have most often been tested using male contests for mates, territories or food. We studied female contests for nest sites and mates in the pied flycatcher, Ficedula hypoleuca, and asked which factors affect the outcome and duration of the contests. Conflicts over residency were experimentally induced, but natural cases in which both females regarded themselves as owners were also included in the analyses. The ensuing escalated contests involved physical fighting, defence of the nestbox of the male and alarm calling. Contrary to expectations from game theory, fighting seemed to occur most often at the start of conflicts, whereas alarm calling occurred towards the end and nestbox visits throughout the conflict. The outcome of the contests was not determined by asymmetries in body size or age, nor by a simple previous-present owner asymmetry. Instead, it depended on the relative residence times of the opponents. The duration of the contests tended to increase with decreasing asymmetry in the residence times of females, whereas body size asymmetries had no influence. Because there is intense competition between females for a mate in the pied flycatcher, we suggest that females do not respect asymmetries in residency and body size, but fight in relation to the value of the mating opportunity. In particular, we point to the possibility that the value of a mating opportunity may increase with residence time because knowledge of other mating options may become outdated.     

The evolution of respect for property

Although possession is ‘nine‐tenths of the law’, respect for ownership is widespread in the animal kingdom even without third‐party enforcement. Thus, the first individuals to find objects are frequently left unchallenged by potential competitors and tend to win contests when disputes arise. Game theory has shown that respect for ownership (‘Bourgeois’ behaviour) can arise as an arbitrary convention to avoid costly disputes. However, the same theory predicts that a paradoxical respect for lack of ownership (‘anti‐Bourgeois’ behaviour) can evolve under the same conditions and in some cases is the only stable outcome. Despite these predictions, anti‐Bourgeois behaviour is rare in nature, whereas respect for ownership is frequently not absolute. Here, we review extensions of the classic models involving repeated interactions, confusion over roles, strategic coordination of behaviour (‘secret handshakes’), owner–intruder asymmetries and continuous control of fighting investment. Confusion over roles and owner–intruder asymmetries in fighting ability may explain why respect for ownership is often partial. Moreover, although most model extensions facilitate the evolution of Bourgeois‐like behaviour, secret handshakes and continuous control of fighting investment render the alternative anti‐Bourgeois convention unstable. We develop these insights to highlight several key areas for future investigation.     

Testing game theory models: fighting ability and decision rules in chameleon contests

Game theory models of animal contests make many non-mutually exclusive predictions, complicating empirical tests. These predictions regard the relationship between contest parameters and fighting ability, for which body size is usually used as a proxy. However, in many systems, body size may be a limited proxy since multiple traits and contextual factors such as experience influence fighting ability. Using contests between male Cape dwarf chameleons, Bradypodion pumilum, I test alternative game theory models of extended contests. I show how the most likely candidate model can be identified through a process of elimination, based on tests of key predictions. In addition, I present a measure of fighting ability based on multiple traits that allows ability to change as experience changes. In dwarf chameleons, persistence is based on loser thresholds rather than assessment of relative ability, ruling out the sequential assessment model. Winners and losers do not match behaviours in early parts of the contest, arguing against all types of war of attrition models. Although the cumulative assessment model remained as the most likely candidate model, not all specific predictions of this model were upheld.     

Combat and communication in the Everglades pygmy sunfish

Changes in population density and resource patterning affect the aggressive behaviour of pygmy sunfish (Elassoma everglaidei) in many ways. Increases in density significantly reduced the proportion of fights directly over clumped prey, the likelihood of initiators winning contests or acquiring clumped prey, and the length of contests. Increases in prey dispersion also reduce the proportion of resource fights in low-and high-density populations, the greater tendency for subordinates to initiate fights directly over resources, the likelihood of initiators acquiring a contested prey item in lowdensity populations, and the length of contests. Such increases, however, increase the likelihood of initiators winning contests at moderate competitive levels, and increase the effectiveness of rapid sequential communication in populations that abandon territoriality. In addition, some of these findings, such as the inverse relationship between contest length and both prey dispersion and rank differential, are consistent with predictions of cost-benefit models of fighting behaviour, namely, that escalated contests become more likely as asymmetries in fighting ability decrease and asymmetries in resource valuation simultaneously increase.     

The assessment game in sand fiddler crab contests for breeding burrows

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

Startle durations reveal visual assessment abilities during contests between convict cichlids

In animal contests, the widespread ability of contestants to assess their opponents’ resource holding potential (RHP) relative to their own, termed mutual assessment, has recently been questioned. It is possible that each contestant may only have information about its own abilities or state, incurring costs up to a particular threshold then giving up, termed self-assessment. We used a technique that provides a measure of fight motivation to discriminate between different assessment models during aggressive encounters between male convict cichlids, Amatitlania nigrofasciata. A novel stimulus was applied to cause a startle response in one contestant of an aggressively interacting, size mis-matched pair, whereby the animal temporarily stops fighting. The time taken to resume the contest has been verified to provide a measure of the motivation to fight, from which it is possible to infer if any visual information concerning opponent asymmetries has been gathered. The data showed support for two differing types of assessment. There was some support for self-assessment, with startle duration being negatively related to own size, and, in a later trial series some support for an opponent only assessment strategy, with startle duration being positively related to opponent size. These results are consistent with individuals learning to use visual information about opponents when deprived of other sensory cues. Evidence within a trial supporting visual mutual assessment was lacking and possible reasons for this are discussed.     

Is Contest Experience a Trump Card? The Interaction of Residency Status, Experience, and Body Size on Fighting Success in Misumenoides formosipes (Araneae: Thomisidae)

Game theory models predict that individuals in contests adjust their strategy appropriately to the current value of the contested resource and the resource holding potential (RHP) of their opponent. In the current investigation, I examined interactively operating, multiple contest asymmetries on dyadic disputes for precopulatory guarding positions in the crab spider Misumenoides formosipes. In contests between equally sized adult males with no previous contest experience, residents had clear advantages in fighting success over intruders. Asymmetries in experience predicted outcome when tested against residency status, and experience operating in concert with residency status predicted resolution when tested against size asymmetries. Data from this investigation suggest that crab spiders learn strategies through experience rather than rely solely on the assessment of their opponent's RHP before determining contest effort.     

Familiarity influences body darkening in territorial disputes between juvenile salmon

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

The importance of being gravid: egg load and contest outcome in a parasitoid wasp

Owners have often been found to have an advantage in animal contests. One explanation of this is that the resource under dispute is of greater value to the owner than to the intruder (nonowner). Such 'resource value asymmetries' may be caused by intrinsic factors, such as the physiological state of the contestants. Females of the bethylid wasp Goniozus nephantidis fight for possession of paralysed hosts. A recent study found that larger females tend to win but owner status also confers an advantage. We tested the hypothesis that ownership advantage is linked to differences in egg load (the number of unlaid ripe eggs) between contestant females. Egg loads were highly variable and positively influenced by age, nutrition, body size and host ownership. Owners almost always won owner-intruder contests. The outcomes of owner-owner contests (in which size and status asymmetries were absent) were strongly influenced by differences in egg load. Copyright 2000 The Association for the Study of Animal Behaviour.     

Using the mixed model for interval mapping of quantitative trait loci in outbred line crosses

Interval mapping by simple regression is a powerful method for the detection of quantitative trait loci (QTLs) in line crosses such as F2 populations. Due to the ease of computation of the regression approach, relatively complex models with multiple fixed effects, interactions between QTLs or between QTLs and fixed effects can easily be accommodated. However, polygenic effects, which are not targeted in QTL analysis, cannot be treated as random effects in a least squares analysis. In a cross between true inbred lines this is of no consequence, as the polygenic effect contributes just to the residual variance. In a cross between outbred lines, however, if a trait has high polygenic heritability, the additive polygenic effect has a large influence on variation in the population. Here we extend the fixed model for the regression interval mapping method to a mixed model using an animal model. This makes it possible to use not only the observations from progeny (e.g. F2), but also those from the parents (F1) to evaluate QTLs and polygenic effects. We show how the animal model using parental observations can be applied to an outbred cross and so increase the power and accuracy of QTL analysis. Three estimation methods, i.e. regression and an animal model either with or without parental observations, are applied to simulated data. The animal model using parental observations is shown to have advantages in estimating QTL position and additive genotypic value, especially when the polygenic heritability is large and the number of progeny per parent is small.     

A TEST OF THE SEQUENTIAL ASSESSMENT GAME: FIGHTING IN THE BOWL AND DOILY SPIDER FRONTINELLA PYRAMITELA

Male bowl and doily spiders (Frontinella pyramitela: Linyphiidae) engage in dangerous fights over access to females. Relatively smaller individuals are more at risk of fatal injury than their larger opponents. Males assess relative fighting ability during contests: smaller individuals tend to give up quickly. Fights occur between a male with information about the value of the contested female (number of fertilizable eggs) and an intruding male with less information. In this paper, a sequential assessment game (a game theory model of fighting behavior) is adapted to male combat in the bowl and doily spider to attempt a quantitative test. The model makes predictions about fight duration, probability of winning, and the occurrence of fatalities as a function of resource value and size asymmetry. Comparison with empirical data from staged contests yields a generally good quantitative agreement with the predictions. A few deviations are also noted.     

What Determines the Duration of War? Insights from Assessment Strategies in Animal Contests

Interstate wars and animal contests both involve disputed resources, restraint and giving up decisions. In both cases it seems illogical for the weaker side to persist in the conflict if it will eventually lose. In the case of animal contests analyses of the links between opponent power and contest duration have provided insights into what sources of information are available to fighting animals. I outline the theory of information use during animal contests and describe a statistical framework that has been used to distinguish between two strategies that individuals use to decide whether to persist or quit. I then apply this framework to the analysis of interstate wars. War duration increases with the power of winners and losers. These patterns provide no support for the idea that wars are settled on the basis of mutual assessment of capabilities but indicate that settlement is based on attrition. In contrast to most animal contests, war duration is as closely linked to the power of the winning side as to that of the losing side. Overall, this analysis highlights a number of similarities between animal contests and interstate war, indicating that both could be investigated using similar conceptual frameworks.     

On Extrapolating Past the Range of Observed Data When Making Statistical Predictions in Ecology

Ecologists are increasingly using statistical models to predict animal abundance and occurrence in unsampled locations. The reliability of such predictions depends on a number of factors, including sample size, how far prediction locations are from the observed data, and similarity of predictive covariates in locations where data are gathered to locations where predictions are desired. In this paper, we propose extending Cook’s notion of an independent variable hull (IVH), developed originally for application with linear regression models, to generalized regression models as a way to help assess the potential reliability of predictions in unsampled areas. Predictions occurring inside the generalized independent variable hull (gIVH) can be regarded as interpolations, while predictions occurring outside the gIVH can be regarded as extrapolations worthy of additional investigation or skepticism. We conduct a simulation study to demonstrate the usefulness of this metric for limiting the scope of spatial inference when conducting model-based abundance estimation from survey counts. In this case, limiting inference to the gIVH substantially reduces bias, especially when survey designs are spatially imbalanced. We also demonstrate the utility of the gIVH in diagnosing problematic extrapolations when estimating the relative abundance of ribbon seals in the Bering Sea as a function of predictive covariates. We suggest that ecologists routinely use diagnostics such as the gIVH to help gauge the reliability of predictions from statistical models (such as generalized linear, generalized additive, and spatio-temporal regression models).     

Visual information, resource value, and sequential assessment in convict cichlid (Cichlasoma nigrofasciatum) contests

We staged contests between convict cichlids (Cichlasoma nigrofasciatum)that were matched for size and gender to test the influenceof prior information and resource value on the duration andstructure of fights. The contestants were separated before thecontest by either clear or opaque dividers to allow or preventvisual assessment, respectively. Contests were shorter in the"clear" than in the "opaque" treatment, suggesting that visualassessment occurred. The duration of lateral display, a noncontactdisplay, was shorter in the clear than in the opaque treatment,but the treatments did not differ significantly in the durationof three contact displays (biting, mouth wrestling, and circling).These results are consistent with the hypothesis that lateraldisplay provides primarily visual information, probably aboutbody size, whereas the other behavior patterns provide primarilynonvisual information, probably about strength. Second contestsbetween the same pair of fish were shorter than first contests,suggesting that the information acquired during the first contestmade it easier to resolve the second. After the subordinatefish from the second contest was given access to a mate, itfought more persistently so that third contests were longerthan second contests. Our results support the predictions ofthe sequential assessment model.     

Asymmetric Forceps Increase Fighting Success among Males of Similar size in the Maritime Earwig

Extreme asymmetric morphologies are hypothesized to serve an adaptive function that counteracts sexual selection for symmetry. However, direct tests of function for asymmetries are lacking, particularly in the context of animal weapons. The weapon of the maritime earwig, Anisolabis maritima, exhibits sizeable variation in the extent of directional asymmetry within and across body sizes, making it an ideal candidate for investigating the function of asymmetry. In this study, we characterized the extent of weapon asymmetry, characterized the manner in which asymmetric weapons are used in contests, staged dyadic contests between males of different size classes, and analyzed the correlates of fighting success. In contests between large males, larger individuals won more fights and emerged as the dominant male. In contests between small males, however, weapon asymmetry was more influential in predicting overall fighting success than body size. This result reveals an advantage of asymmetric weaponry among males that are below the mean size in the population. A forceps manipulation experiment suggests that asymmetry may be an indirect correlate of a morphologically independent factor that affects fighting ability.     

Comparison of Logistic Regression and Linear Regression in Modeling Percentage Data

Percentage is widely used to describe different results in food microbiology, e.g., probability of microbial growth, percent inactivated, and percent of positive samples. Four sets of percentage data, percent-growth-positive, germination extent, probability for one cell to grow, and maximum fraction of positive tubes, were obtained from our own experiments and the literature. These data were modeled using linear and logistic regression. Five methods were used to compare the goodness of fit of the two models: percentage of predictions closer to observations, range of the differences (predicted value minus observed value), deviation of the model, linear regression between the observed and predicted values, and bias and accuracy factors. Logistic regression was a better predictor of at least 78% of the observations in all four data sets. In all cases, the deviation of logistic models was much smaller. The linear correlation between observations and logistic predictions was always stronger. Validation (accomplished using part of one data set) also demonstrated that the logistic model was more accurate in predicting new data points. Bias and accuracy factors were found to be less informative when evaluating models developed for percentage data, since neither of these indices can compare predictions at zero. Model simplification for the logistic model was demonstrated with one data set. The simplified model was as powerful in making predictions as the full linear model, and it also gave clearer insight in determining the key experimental factors.     

All by myself? Meta‐analysis of animal contests shows stronger support for self than for mutual assessment models

Since the 1970's, models based on evolutionary game theory, such as war of attrition (WOA), energetic war of attrition (E‐WOA), cumulative assessment model (CAM) and sequential assessment model (SAM), have been widely applied to understand how animals settle contests. Despite the important theoretical advances provided by these models, empirical evidence indicates that rules adopted by animals to settle contests vary among species. This stimulated recent discussions about the generality and applicability of models of contest. A meta‐analysis may be helpful to answer questions such as: (i) is there a common contest rule to settle contests; (ii) do contest characteristics, such as the occurrence of physical contact during the fight, influence the use of specific contest rules; and (iii) is there a phylogenetic signal behind contest rules? To answer these questions, we gathered information on the relationship between contest duration and traits linked to contestants' resource holding potential (RHP) for randomly paired rivals and RHP‐matched rivals. We also gathered behavioural data about contest escalation and RHP asymmetry. In contests between randomly paired rivals, we found a positive relationship between contest duration and loser RHP but did not find any pattern for winners. We also found a low phylogenetic signal and a similar response for species that fight with and without physical contact. In RHP‐matched rivals, we found a positive relationship between contest duration and the mean RHP of the pair. Finally, we found a negative relation between contest escalation and RHP asymmetry, even though it was more variable than the other results. Our results thus indicate that rivals settle contests following the rules predicted by WOA and E‐WOA in most species. However, we also found inconsistencies between the behaviours exhibited during contests and the assumptions of WOA models in most species. We discuss additional (and relatively untested) theoretical possibilities that may be explored to resolve the existing inconsistencies.     

Individual-based simulations of the War of Attrition

The War of Attrition model of John Maynard Smith predicts a single, mixed evolutionarily stable strategy (ESS) for animal contests which are settled by conventional displays with no assessment of the opponent's fighting ability. We test the predictions of the model by simulating the evolution of strategies in a finite population of animals under various assumptions on how possible strategies are coded and mutated. While our simulations for the most part confirm the predictions of the model, we also discovered some significant deviations from the theoretically predicted ESS. Specifically, we found that if inheritance of strategies is somewhat imprecise, then a population can evolve that achieves on average a higher payoff than a population at the theoretically predicted ESS. Moreover, if the ESS is realized as a polymorphism of fixed persistence times, then for small populations, sufficiently stringent statistical tests will reject the hypothesis that these times are distributed as theoretically predicted.     

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

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

Individual participation in intergroup contests is mediated by numerical assessment strategies in black howler and tufted capuchin monkeys

Asymmetries in resource-holding potential between opposing groups frequently determine outcomes of intergroup contests. Since both numerical superiority and high intergroup dominance rank may confer competitive advantages, group members should benefit from assessing the relative strength of rivals prior to engaging in defensive displays. However, differences in individual assessment may emerge when cost–benefit trade-offs differ among group members. We examine the influence of numerical superiority and intergroup dominance relationships on individual participation in intergroup encounters in black howler monkeys (Alouatta pigra) and tufted capuchin monkeys (Sapajus nigritus). Black howlers responded with longer vocal displays during encounters with neighbours with an equal number of resident males, while tufted capuchins increased their participation with increasing relative male group size. Within each species, males and females responded similarly to varying numerical odds, suggesting that despite pay-off asymmetries between males and females, both sexes were similarly influenced by numerical asymmetries in deciding to participate in collective group defence. Whereas the outcome of contests among tufted capuchins was determined by relative male group size, reflected in a pronounced intergroup dominance hierarchy, the absence of dominance relationships among black howler groups may have provoked prolonged vocal displays in order to assess rival groups with matching competitive abilities.