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.     

Does prior residency interact with loss? A study of male–male contests in the hermit crab Pagurus minutus

Residency is an important predictor of success in contests with ownership asymmetries. Residency often can interact with a winning experience. However, given that some residents lose a contest even when showing an ownership advantage and that the process leading to loss often determines the loser's subsequent success, prior ownership might also interact with a loss. Here, we staged experimental contests between males of the hermit crab Pagurus minutus with a similar-sized weapon (i.e., cheliped) to examine this possibility. Male–male contests in this species occur between a solitary intruder and an owner guarding a mature female. We evaluated (a) whether resource ownership and female value affect the contest outcome and (b) whether the probability of winning after losing differs depending on the initial role of the loser (i.e., owner or intruder) by using precopulatory guarding pairs of P. minutus collected from the field. In the first fighting trial, we found an ownership advantage and increasing owner success as the body size of his partner increased. Although some owners lost the fight, in contrast to our prediction, the frequency of losing in the second fighting trial did not differ between prior owners and prior intruders. Because losers from the first fighting trial of male–male contests have no female regardless of their initial role, this shared solitary status might be related to the lack of difference in success in the second fighting trial. Moreover, unlike in other animals, resident status might not always assure greater fighting ability in P. minutus males because guarding Pagurus males can avoid male–male contests by climbing up objects in the field. Losers in the first trial, therefore, may have been weaker contestants based on traits other than size, regardless of whether they were owners or intruders.     

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.     

Power to Hold Sheltering Burrows by Juveniles of the Signal Crayfish,Pasifastacus leniusculus

Juveniles of the signal crayfish reside during daylight hours in shelters. At twilight they leave for food, at daybreak they either return or find another shelter. We examined conflicts over burrow ownership. At low densities with equal numbers of shelters and crayfish 1/5 of the burrows were occupied. Increasing both the number of crayfish and shelters improved the occupancy close to 50%. Doubling the number of crayfish in relation to the number of shelters increased the occupancy up to 75%. In an experiment with 30 randomly selected crayfish and 15 sheltering holes available the burrow holders were about 1–2 mm longer in carapace length than those found freely moving in the aquarium. Shelter owners were rarely newly moulted individuals. Size asymmetry (2 mm difference in carapace length) between owner and intruder affected the outcome of the contest over burrow ownership. When intruders were larger than owners, takeovers occurred in about 80% of the cases tested. If the owner was large and the intruder small the takeover frequency was about 20%. When the owner and the intruder were of matching size takeovers were still observed in about 45% of the cases. The outcome of the ownership contest has a true meaning. In a nursery-pond experiment, where low-protein food was randomly scattered all around, no size differences were found in carapace lengths of crayfish juveniles residing in different shelters. However, when high-protein food was introduced in a single spot, owners of burrows were significantly larger and more numerous in the nearby shelters than in the shelters furthest away from the food source. Therefore, when food is unevenly distributed the burrow ownership contests may potentially lead to size asymmetries between individuals. This may lead to large individuals nearby the food growing faster than small individuals further away from the food source, a process likely to further enhance size differences.     

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.

     

Early Comers Become Owners: Effect of Residency Experience on Territorial Contest Dynamics in a Lycaenid Butterfly

A positive relationship between ownership and contest success is widespread among territorial butterflies. We tested the hypothesis that owners win simply because they have a longer experience occupying a given territory than do intruders in territorial butterflies. Using the lycaenid butterfly Chrysozephyrus smaragdinus , we staged contests between two males that had experienced different residency periods at the same contested territory, similar to owners and intruders in natural contests. We found that males that had occupied the contested territory for a longer period (a day) always defeated the shorter occupancy males (15 min). Therefore, residency period plays a key role in settling territorial contests. As residency confers advantages in a time-dependent manner, the mechanism of owner dominance could be an increase in motivation to fight as local familiarity increases. Moreover, this result predicts a population-level territorial dynamics. If residency period per se confers advantages in each contest, males arriving at a territory early in their adult season should acquire territorial dominance because they can remain there for a certain period with only a few rivals; late-appearing males do not have this advantage. As predicted, we found that males appearing at territory sites early in the season were more likely to hold their territories. This finding provides new evidence for the evolution of protandry in butterflies.     

Residency Duration and Shelter Quality Influence Vibratory Signalling Displays in A Territorial Caterpillar

Residents are more likely to win territorial disputes than intruders. One explanation for this prior resident advantage is that residents place a higher value on the resource and are therefore more motivated to win. Although value asymmetry models of animal contests often assume that contestants use information about resource value, information on the proximate cues affecting territorial behaviour is often lacking. We use a simple model system – territorial behaviour in the masked birch caterpillar (Drepana arcuata) to identify factors that affect territorial behaviour. Late instar caterpillars occupy solitary silken leaf shelters, which they defend against wandering conspecifics with a vibratory display. We evaluated how a caterpillar identifies itself as the owner and the factors that influence a resident's motivation to signal. To do so, we conducted three experiments between size‐matched residents and intruders to assess how residency duration and shelter quality independently affected territorial displays during the early stages of a contest. Experiment 1 (Time Exp.) demonstrated that resident signalling rates increase with increased duration on the leaf prior to introducing the intruder. Residents also signal more than intruders after residency periods of 1–3 min and longer, demonstrating that residents gather information about resource value shortly after occupying a leaf. Experiment 2 (Squatter Exp.) aimed to disentangle the effects of time on the leaf and silk accumulation. Squatters (individuals in a shelter made by another) placed for 1–3 min on a leaf containing a full silk shelter signalled more to intruders than did caterpillars placed on a fresh leaf for 1–3 min. Experiment 3 (Shelter Removal Exp.) showed that residents whose shelters had been removed signal less than those occupying an intact shelter, despite an equal length of time investing in them. Our experiment is the first to covary both prior residency duration and territory quality, and we find that the motivation of caterpillars to signal is a function of both of these attributes.     

Assessment of fish size on shelter choice and intraspecific interactions by round gobies <Emphasis Type="Italic">Neogobius melanostomus</Emphasis>

Synopsis We examined shelter occupancy and behavioural interactions in non-reproductive male round gobies, Neogobius melanostomus,an invasive fish, to determine if gobies can assess one another’s prowess effectively. Results of laboratory experiments revealed a significant, positive relationship between body size (total mass) and time for individual fish to occupy shelters. Shelter size selection did not vary with body size, but males that abandoned shelters were smaller than those that remained. Overall, the number of interactions between residents and intruders were low with most interactions occurring between large residents and large intruders. A size differential between residents and intruders of 3% was sufficient to predict the proportion of victories in conflicts between round gobies. This ability of the round goby to perceive size differences between themselves and potential opponents reduces the number of interactions and may account for the occurrence of high densities of round gobies observed in the field.     

Sexual dominance in hermit crab shell fights: asymmetries in owner–intruder status,crab size,and resource value between sexes

We studied sexual dominance and seasonal differences in aggressiveness of individuals in intraspecific competition for shells of the hermit crab Pagurus filholi in terms of size of contestants and duration of the attempt to deprive other crabs of their shell. Experiments were conducted using paired intrasexual and intersexual contests in the pre-breeding and post-breeding seasons. Size ratios between contestants were systematically varied to assess the sexual difference in size and owner advantages. In both intrasexual and intersexual contests intruder crabs tended to win the contests more often as their size increased, that is, size advantage overcame owner advantage. Although we did not recognize a sexual difference in size and owner advantages in contest outcomes, male intruder crabs took a shorter time to deprive female owners of a shell than to deprive male owners. Furthermore, male individuals in the pre-breeding season had significantly longer fight durations. Fighting is costly. Thus males can afford to expend more energy and time fighting, indicating that males are dominant over females in shell fights as both intruders and owners. Electronic Publication     

Role assessment,reserve strategy,and acquisition of information in asymmetric animal conflicts

It was formerly argued that alternative evolutionarily stable strategies (ESSs) are possible for animal contests characterized by some asymmetry that can be perceived with perfect accuracy. Where roles A and B refer to the asymmetry between opponents, ESSs are: ‘fight when A, retreat when B’, and vice versa. Either can be an ESS, but only if the ‘reserve strategy’ (=what an animal does when it fights) is sufficiently damaging. We examine the ‘war of attrition’ (winner = opponent that persists longer). In a population at either ESS, reserve strategy is never normally shown; it is therefore subject to drift unless the selective action of rare individuals which break the convention is considered. These could arise either by mutation or by mistakes in role assessment. When mutations and mistakes simply specify that occasionally an animal fights when it ‘should’ retreat, selection adjusts reserve strategy to a level where only one ESS (the ‘commonsense’ ESS) is possible, if the asymmetry is relevant to payoff. Thus for asymmetries in fighting ability or resource value, the individual with the lower score will retreat. However, we are particularly concerned with cases where both payoff-relevant aspects (fighting ability and resource value) are asymmetric. If opponents sustain contest costs at rates K_A and K_B, and their resource values are V_A and V_B, an ‘optimal assessor’ strategy defined by the interaction between the two asymmetries, is a unique ESS. It obeys the rule ‘fight on estimating role A, where V_A/K_A>V_B/K_B; retreat in B’. If mistakes can occur in both roles, but are very rate, the ESS is not fundamentally altered though there will be infinitesimal tendencies for persisting in role B. Selection to improve assessment abilities intensifies as abilities improve, but is weak if roles A and B are rather similar. Over a range of similarity between roles, an ‘owner wins’ convention may be adopted if ownership correlates positively with role A and an individual cannot tell when it would otherwise pay him to break the convention. We also examine a contest in which information about roles can be acquired only during a contest itself, and at a cost. Much depends on the rate at which information is acquired relative to the rate at which costs are expended, and on whether contests normally escalate in intensity, remain at the same level, or de-escalate. Selection favours short contests when costs are high relative to resource value, where the outcome of a round contains much information about fighting ability, and where the actual disparity in fighting ability is large.     

Influence of asymmetries in resource quality and age on aggression and dominance in black-chinned hummingbirds

According to current theory, dominance relationships may depend on asymmetries between adversaries in their expected net gain from contested resources. A prediction derived from this theory was confirmed in the field by experimentally creating asymmetries between neighbouring territorial hummingbirds with regard to their net energy gains from their territories, and then forcing them to fight for ownership of the combined resources of the two territories. When territory productivity was increased to levels well above the daily energy requirements of each contestant, energetic asymmetries no longer detectably influenced dominance, and the amounts of aggression per contest were lower. Contests in which an energetic asymmetry was partially offset by an age-related asymmetry involved more fights than contests in which opponents belonged to different age classes but controlled feeders of identical productivity. The predicted negative correlation between the magnitude of the asymmetry in net energy gain and agonistic investment in the contest did not occur.     

Field Observations of Agonism in the Crayfish, Orconectes rusticus: Shelter Use in a Natural Environment

Crayfish are aggressive animals that compete to acquire resources such as shelters, food, and mates. Shelters are a primary resource that crayfish use for protection from conspecifics and predators. Despite the importance of this resource, no field research has been performed that studies the acquisition and control of this resource. The present study examines shelter use in a natural habitat and the impact that shelter ownership has on the intensity and outcome of agonistic encounters. A stationary underwater camera was used to observe crayfish, Orconectes rusticus, shelter use and agonistic interactions in a natural lake environment. These shelters were formed naturally in iron outcroppings found on the limestone and detritus benthos. Crayfish activity and shelter use was found to be dependent upon a circadian cycle with most of the shelter use occurring during the morning to early afternoon (05:00–13:00 hours). Agonistic encounters in the presence of shelters resulted in short, low intensity interactions. Interestingly, fight outcomes were not significantly affected by shelter ownership, but were primarily determined by size differential of combatants. This outcome may be due to the prevalence of shelters within this habitat. Contrary to laboratory studies, these results indicate that shelter ownership may not be an important factor in determining resource‐holding potential in some habitats.     

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.     

Matter of Size or Matter of Residency Experience? Territorial Contest in a Green Hairstreak,Chrysozephyrus smaragdinus (Lepidoptera: Lycaenidae)

Territorial residents usually win asymmetrical owner‐intruder contests and a variety of hypotheses have been proposed to explain this phenomenon. In the butterfly Chrysozephyrus smaragdinus (Lepidoptera: Lycaenidae), male territorial residents defended their territories against intruders during numerous contests and kept them for many successive days. Field observations and experiments were conducted to examine the factors related to this superiority of residents. Forewing length did not differ between residents and intruders, suggesting that body size is not correlated with resource holding potential. Removal–replacement experiments demonstrated that residency did not serve as an arbitrary means for contest settlement, and did not support the recently presented alternative hypothesis that males with higher body temperature are more likely to win. New residents fought longer in defense of the territory as their residence duration in the territory increased. I discuss these results in light of game theory and suggest that the superiority of residents in C. smaragdinus may be based on the asymmetry of resource (territory) value for residents and intruders.     

Fighting in Males of the Autumn Spider,Metellina segmentata: Effects of Relative Body Size,Prior Residency and Female Value on Contest Outcome and Duration

Males of the autumn spider, Metellina segmentata (***Araneac: Metidae), compete for access to mates by guarding the orb webs of mature females. We investigated the influences of relative male righting ability and resource value on fighting behaviour by staging interactions in the field on webs occupied by females. In these contests, the larger male nearly always defeated its opponent when it was at least 10% greater in size. For smaller size asymmetries between opponents, the male previously resident on the female's web usually won the contest. Contest duration decreased exponentially with increasing size asymmetry between opponents, as predicted if each male assessed its relative size and adjusted its fighting strategy according to its likelihood of winning. Contest duration was also greater when the prior resident was the lighter opponent, or when size and residency asymmetries favoured different opponents as winners. Prior residents fought longer over larger, more fecund females, indicating an adjustment of fighting effort according to assessments of resource value. In contrast, intruders did not increase their fighting effort over larger females, suggesting an inability to assess female size quickly and accurately. Assessment appears to reduce the costs of settling conflicts, but imperfect information can result in inaccurate assessments and unexpected outcomes. Assessment strategies are used by other types of spiders to resolve contests, but this appears to be the first evidence for such strategies among orb-web-building spiders.     

Shelter Availability,Occupancy, and Residency in Size‐Asymmetric Contests Between Rusty Crayfish,Orconectes rusticus

Contest outcomes are usually determined by differences in resource‐holding potential, the social histories of the combatants, and perceptions of resource value. One aspect of gaining an advantage is the residency effect. Prior occupancy of a particular place can affect the knowledge and motivation of the resident. There could be a tactical advantage in knowing the terrain or an increased willingness to fight to maintain control of a familiar area. In this study, we evaluated the importance of shelter residency effects relative to size differences between rusty crayfish (Orconectes rusticus) as potential competitors for access to shelter. The intensity of any residency effects was manipulated by altering the number of shelters in the arena. Our results suggest that any residency effect is very weak in this system, and if present, may often be masked by the strong and pervasive influence on contest outcome of the relative body sizes of the contestants. We also found that both shelter number and crayfish size asymmetries had strong, independent effects on levels of aggression. Dominance, but not residency status, was a factor in shelter use.     

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.     

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.     

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.