Ownership,size and reproductive status affect the outcome of food ball contests in a dung roller beetle: when do enemies share?

Theory predicts that asymmetry between contenders influences their ability to defend resources. More recently, some theoretical approaches have also examined the circumstances that might promote sharing of the disputed resources. We tested these hypotheses in males of the ball roller beetle Canthon cyanellus cyanellus. Males fight for possession of a food ball, which is a vital resource used for nesting. We evaluated the role of food resource ownership, body size and reproductive status on the outcome of contests (win, lose or share) between males that rolled a food ball (owners or finders) either alone or with a female partner, when faced with male intruders (or joiners). Large owners of a food ball had a higher probability of victory than small intruders, and small owners had a high probability of losing when faced with large intruders. The reproductive status of both contenders also influenced their chances of winning: previously mated owners of a food ball had a higher probability of winning than virgin owners. Males of a similar size tended to split the food ball, thereby sharing the resource. Our results suggest that competitors may adjust the intensity of their aggression depending at least on their own resource holding power (RHP), the value of the resource in dispute and perhaps even the RHP of their opponents. Sharing the food ball emerges as a fresh solution between similarly matched contestants.     

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

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

The relation between mate value, entitlement, physical aggression, size and strength among a sample of young Indian men

This study extends previous ones showing a link between direct aggression and size and strength among young men, which were informed by the evolutionary concept of resource holding power (RHP), using measures of size, strength, flexed bicep circumference and hand grip strength among a sample of young men from the Indian state of Mizoram. The study also examined the relation of these variables to reactive and proactive aggression, to entitlement to resources (related to the threatened egotism theory of aggression) and mate value (central to a modular theory of self-esteem and more broadly to sexual selection). The findings showed only a weak association between size and strength and direct aggression, which was also significantly correlated with entitlement and mate value, as predicted. Mate value also showed some association with size and strength. Reactive but not proactive aggression was linked with entitlement, but neither was associated with size or strength. In a regression analysis, controlling for age, mate value was the strongest predictor of direct aggression, but both weight (highly correlated with strength) and entitlement were marginally significant predictors independent of mate value. The findings provide support for the view that young males who view themselves as more attractive to women are more aggressive, independently of the impact on aggressiveness of RHP or entitlement to resources.     

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.     

Intruder Pressure Affects Territory Size and Foraging Success in Asymmetric Contests in the Water Strider Gerris lacustris

The theory of evolutionarily stable strategies (ESS) in asymmetric contests predicts that the propensity of an individual to expose itself to risk during contests depends on the individual's resource-holding-potential (RHP) and on the value of the disputed resource (V) for the individual compared with that for an opponent. If encounters of a territory owner with individuals of high RHP and high food demands (V) increase in frequency, one should expect a decrease in total aggressiveness of the territory owner, and in consequence a decrease of its territory size. Such a decrease should result in a lower amount of food consumed by the territory owner. Using natural variability in RHP and V in Gerris lacustris, I experimentally tested these predictions. The average prey item has higher value (V) for reproductive female water striders (which probably transform most of their food into eggs), than for nonreproductive females and for males. Because males are smaller, they have lower RHP than females, as RHP depends on size. Thus the reproductive females are the class of individuals of high RHP and high food demands (high V). Most nonreproductive females defend food-based territories. I observed two groups of water striders in a seminatural laboratory setting. As predicted, there was a negative correlation between the rate of encounter with reproductive females and size of the territory, and a positive correlation between territory size and number of Drosophila flies consumed by the owner. Territories were smaller in the group with high rates of encounter between territory owners and reproductive females. Territory owners caught the same number of Drosophila flies as non-territorial individuals in this group. In contrast, in the group with fewer encounters between territory owners and reproductive females, territories were larger, and territory owners gained more food than non territorial water striders.     

Territory size in the moorhen (Gallinula chloropus): An outcome of RHP asymmetry between neighbours

Territory size can be considered as the outcome of contests between an owner and its neighbours for the resource units in the defended area. Variation in territory size could be dependent upon three possible asymmetries: (1) difference in resource holding potential (RHP); (2) difference in resource value to the competitors; and (3) difference in ‘ownership’ status. These possibilities were tested in a 3-year study of moorhens which defend linear territories along ditches in a grazing marsh habitat. Of the three, relative difference in RHP between an owner and its two neighbours provided the best correlation with territory size. There was no relationship between the amount of resources per unit area and territory size nor between the time of establishing a territory and its later size. Heavier moorhens are more likely to win contests in winter flocks and are thought to have a greater RHP. The ratio of the weight of a male territory owner to the weight of both its male neighbours was highly correlated with territory size. Birds with larger territories may gain more potential nesting sites with good cover from predators.     

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.     

Effect of environmental enrichment upon resource holding power in fish in prior residence situations

Resource holding power (RHP), as expressed by gaining dominance, can be affected by extrinsic and intrinsic factors. Extrinsic factors that increase the RHP include e.g. prior exposure to the contest area. The pay-off asymmetry hypothesis was tested according to which there is an asymmetry in the value the resident's territory has for the resident and intruders, i.e. the resident loses more than intruder opponents when losing the contest over a territory in which it has invested much energy during the exploration of food and shelter resources, during the settlement of conflicts with territorial neighbours, etc. This asymmetry will increase the resident's chance to win the fight over intruders. If the resident-intruder asymmetry is enhanced by presenting resource-rich territories to e.g. fish (structurally diverse or 'rich' aquariums), the hypothesis predicts that the resident's dominance chance (probability to win the fight) will be higher than in structurally 'poor' aquariums. It was found that residents were proportionately more often dominant (fight winners) than intruders in the rich compared to the poor aquariums in all seven tested fish species. This demonstrates that a high territory resource value (aquarium enrichment) significantly facilitates the expression of the resident's dominance advantage (prior residence effect). In contrast to dominance, aggressive behaviour before the dominance settlement did not generally differ between residents and intruders when compared in rich and poor aquariums. This suggests that dominance in itself and aggression prior to dominance settlement are at least partially guided by different motivational systems.     

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.     

Pure Self‐Assessment of Size During Male–Male Contests in the Parasitoid Wasp Nasonia vitripennis

During contest competition, a competitor may persist in a given contest based on information regarding its own fighting ability (resource‐holding potential, RHP), or that of its opponent. Although a number of models formalize the ways in which competitors are hypothesized to use RHP‐related information to determine their persistence in contests, we focused on pure self‐assessment and mutual assessment models in this study. According to pure self‐assessment models, a competitor uses only information regarding its own RHP to determine its persistence in a contest. In contrast, according to mutual assessment models, persistence is based on information regarding a competitor's RHP relative to that of its opponent and therefore requires assessment between competitors. In this study, using size as a proxy for RHP, we tested whether the parasitoid wasp Nasonia vitripennis utilizes pure self‐assessment or mutual assessment during pairwise, male–male contests. When we examined competitors of varied sizes, we found that the losing male's size was positively related to contest duration, but the winning male's size was uncorrelated with contest duration. When we examined contests in which competitors were size‐matched, we found that the mean size of paired competitors was positively related to contest duration. These results suggest that male N. vitripennis engage in pure self‐assessment during contests.     

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.     

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.     

Competition for space can drive the evolution of dormancy in a temporally invariant environment

I present a model for the evolution of a seed bank in the absence of externally driven environmental variation. I use Evolutionarily Stable Strategy (ESS) analyses of both analytic and simulation models to assess the conditions under which a dormant genotype can invade and resist invasion. In my models, plant seeds compete through lottery for discrete safe sites holding one individual each. Analyzing the conditions under which a dormant genotype can invade when rare and resist invasion once established, I conclude that dormancy can be an ESS when some fraction of seeds is retained locally, seed bank survival is high, and mortality in the seed bank is low. The advantage of dormancy stems from the ability of dormant seeds to recapture a lost site and the fact that a plant’s offspring are more likely to win the lottery in its own site than in any new site. The advantage of dormancy does not depend on individual fecundity or on low relatedness with the offspring of kin, making this mechanism distinct from earlier models of sib competition.     

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.     

Subjective and objective components of resource value additively increase aggression in parasitoid contests

Two major categories of factors are predicted to influence behaviour in dyadic contests; differences in the abilities of the contestants to acquire and retain resources (resource holding potential), and the value of the contested resource (resource value, RV; which comprises objective and subjective components). Recent studies indicate that subjective components affect contest behaviour in several animal taxa but few have simultaneously investigated objective RV components. We find that both an objective (host size) and a subjective (contestant age) component of RV affect contest intensity in the parasitoid wasp Goniozus legneri. These additively influence aggressiveness, with a larger effect from the subjective component than the objective component. The greater influence of subjective RV adds weight to the recent surge of recognition of this RV component''s importance in contest behaviour.     

Same resource,different benefits: hermit crab shell structure advantages owners,but not intruders in agonistic interactions

Resources may confer advantages by enhancing their owners’ fighting ability (resource holding potential; RHP). While the resource-correlated RHP hypothesis has been recognized as a determinant of agonistic success in different taxa, this has mostly been based on assessment of either the intruder or the owner, but only rarely in both contestants. We tested whether the internal structure of shells affects hermit crabs’ RHP, both as owners defending the shell against eviction and as intruders attempting to gain access to an occupied shell. We conducted contests (n?=?60) to compare the success in shell eviction by intruders in intact shells vs. in shells with the columella artificially reduced, and the success of shell retention by owners in intact shells vs. shells with the columella reduced. The internal configuration of the shell showed different resource-correlated RHP effects depending on the individual’s role in the fight. The presence of a columella in the intruder’s shell did not affect the likelihood that they would evict their opponents. However, owners resisted more evictions in shells with intact columella than those in shells with reduced columella. Our results demonstrate that the same resource can offer different RHP advantages to owners and intruders during an agonistic interaction.

     

Transitive inference in non-human animals: an empirical and theoretical analysis

Transitive inference has long been considered one of the hallmarks of human deductive reasoning. Recent reports of transitive-like behaviors in non-human animals have prompted a flourishing empirical and theoretical search for the mechanism(s) that may mediate this ability in non-humans. In this paper, I begin by describing the transitive inference tasks customarily used with non-human animals and then review the empirical findings. Transitive inference has been demonstrated in a wide variety of species, and the signature effects that usually accompany transitive inference in humans (the serial position effect and the symbolic distance effect) have also been found in non-humans. I then critically analyze the most prominent models of this ability in non-human animals. Some models are cognitive, proposing for instance that animals use the rules of formal logic or form mental representations of the premises to solve the task, others are based on associative mechanisms such as value transfer and reinforcement and non-reinforcement. Overall, I argue that the reinforcement-based models are in a much better empirical and theoretical position. Hence, transitive inference in non-human animals should be considered a property of reinforcement history rather than of inferential processes. I finalize by shedding some light on some promising lines of research.     

Frustrative reward omission increases aggressive behaviour of inferior fighters

Animals use aggressive behaviour to gain access to resources, and individuals adjust their behaviour relative to resource value and own resource holding potential (RHP). Normally, smaller individuals have inferior fighting abilities compared with larger conspecifics. Affective and cognitive processes can alter contest dynamics, but the interaction between such effects and that of differing RHPs has not been adjudged. We investigated effects of omission of expected reward (OER) on competing individuals with contrasting RHPs. Small and large rainbow trout (Oncorhynchus mykiss) were conditioned to associate a light with reward. Thereafter, the reward was omitted for half of the fish prior to a contest between individuals possessing a 36–40% difference in RHP. Small control individuals displayed submissive behaviour and virtually no aggression. By contrast, small OER individuals were more aggressive, and two out of 11 became socially dominant. Increased aggression in small OER individuals was accompanied by increased serotonin levels in the dorsomedial pallium (proposed amygdala homologue), but no changes in limbic dopamine neurochemistry were observed in OER-exposed individuals. The behavioural and physiological response to OER in fish indicates that frustration is an evolutionarily conserved affective state. Moreover, our results indicate that aggressive motivation to reward unpredictability affects low RHP individuals strongest.     

Intraguild predation in biological control: consideration of multiple resource species

Theoretical models of intraguild predation (IGP) predict that IGP decreases the effectiveness of biological control while many empirical studies do not agree with this prediction. In this study, I discuss the importance of explicit consideration of multiple resource species that has been neglected in most theoretical IGP models. In the previous models of IGP, a single resource species represented the pest species. However, there are multiple resource species (e.g., multiple pest species or aggregates of pest and non-pest species) in real systems. This study shows that models with multiple resource species can predict a variety of outcomes including those consistent with the empirical observations. The explicit consideration of resource species is useful for the future development of theories in biological control. Handling Editor: Helen Roy     

Subjective resource value affects aggressive behavior independently of resource-holding-potential and color morphs in male common wall lizard

Journal of Ethology - Game theory models predict the outcome of a dyadic contest to depend on opponents’ asymmetries in three main traits: resource-holding potential (RHP), resource value...