Fighting without weaponry: a review of male-male contest competition in butterflies

The males of many butterfly species compete via pair-wise intrasexual contests, yet lack any obvious morphological traits conventionally associated with animal aggression. The evolution of contest behavior in this group is therefore interesting because the means by which individuals are able to inflict costs upon each other during combat are unclear. Here we review the relevant empirical information within a game-theoretical context in order to analyze contest-related costs and mechanisms of dispute settlement. Territorial butterfly contests carry obvious role asymmetries, and individuals in the 'resident' role are more likely to win across all studied species. We use this phenomenon as a framework for gaining further insight into the evolution of contest settlement in this group. Four principal hypotheses are evaluated: (1) that residency is used as an arbitrary convention, (2) that residents stand to gain a greater payoff if successful, (3) that residency bestows higher resource-holding potential (RHP) on males in that role, and (4) that residents are males of intrinsically higher RHP. We found most support for the latter explanation (the 'superior-competitor' hypothesis), partly because the alternatives require a restrictive set of ecological conditions, but also due to positive support for the idea of intrinsic RHP. The strongest evidence is provided by species in which repeated interactions (between the same individuals) always have the same outcome, regardless of changes in residency status. However, we also found a consistent effect on contest duration due to apparent 'confusion over residency,' which suggests a conventional element to contest behavior. Although butterfly contests must be costly in some way, the literature provides limited insight into how costs accrue, and hence the determinants of contest RHP remain obscure. We suggest that careful experimentation, using 'natural' manipulations where possible, should prove most beneficial in identifying costs and exploring schedules of contest settlement in this group. Moreover, future investigations should be sensitive to the fact that butterfly territoriality and associated male-male interactions are seated within a broader ecological and life-historical context.     

The role of body size and fighting experience in predicting contest behaviour in the black field cricket, <Emphasis Type="Italic">Teleogryllus commodus</Emphasis>

Body size strongly predicts fighting behaviour and outcome in many species, with the larger opponent usually winning contests. However, recent fighting experience can have a strong influence on the establishment of dominance hierarchies, with recent winners being more likely to win subsequent contests, while recent losers are more likely to lose. Recent fighting experience therefore has the potential to modify the effect of body size in determining contest behaviour and outcome. Here, we investigate whether recent fighting experience weakens the role of body size in predicting contest behaviour in the black field cricket, Teleogryllus commodus. We compared the role of body size in determining contest outcome during initial non-physical encounters and escalated aggressive physical encounters (grapples), as well as the probability of escalation occurring, in contests involving either a naïve or experienced smaller male against a naïve larger male. We found that recent fighting experience only affected contest outcome during non-physical encounters. Once a contest had escalated into grappling, the effect of previous wins and losses was no longer apparent and body size strongly predicted contest outcome. Thus, once males can directly assess their opponent's fighting ability, recent fighting experience did not alter the effect of body size on contest behaviour and outcome.     

Body morphologies shape territorial dominance in the satyrine butterfly <Emphasis Type="Italic">Lethe diana</Emphasis>

Male butterflies compete over mating territories via aerial interactions. It has often been suggested that flight performance, and thus flight-related traits, would play an important role in butterflies’ contest resolution; however, most studies have not demonstrated a clear relationship between these traits and territory ownership in butterflies. Males of the satyrine butterfly Lethe diana compete over territories via linear chases, a behavior that is uncommon among butterflies: rather, contests of most butterfly species consist of circling flights. In the present study, some morphological and physiological traits of L. diana were compared between territory owners and intruders to search for traits correlated with resource holding power (RHP). Territory owners had higher flight-muscle ratio (FMR) and were heavier than intruders, indicating that flight performance plays a key role in their contest resolution. Contest duration was not affected by the owners’ FMR or body mass, and thus I obtained no evidence supporting the hypothesis that contestants assess their relative RHP before making a decision to retreat. I speculate that the fact that these traits had an effect on territorial status in L. diana was a result of the characteristics of the contest behavior of this butterfly. During their contests, males L. diana fly faster in their linear flights than males of most other butterflies, which engage in slower circling flights. Therefore, males of L. diana probably need higher flight performance capabilities in order to win their contests.     

Territorial defence in the great tit (Parus major): Do residents always win?

Summary I removed resident pairs of great tits from their territories for short periods and released them after replacement pairs had occupied the spaces. When two pairs are manipulated in this way into occupying the same territory an escalated contest ensures. Contests between residents and replacements are longer, more likely to involve physical fights and longer fights or displays than contests between established neighbours or residents and intruders. The degree of escalation between residents and replacements is an increasing function of replacement time. It rises to a peak and then diminishes again. The probability that the replacement will defeat the former resident is an increasing function of replacement time, reaching 90% after several days. These results are consistent with the hypothesis that territorial residents win in contests against intruders because of an asymmetry in payoff rather than an asymmetry in resource holding potential or an arbitary convention. A possibly asymmetry in payoff in the great tit is the cost of defending the territory against neighbours. The cost is higher for replacements than for residents.     

Contest behaviour in the speckled wood butterfly (Pararge aegeria): seasonal phenotypic plasticity and the functional significance of flight performance

Although contemporary animal contest theory emphasises the importance of physical asymmetries in resolving disputes, such asymmetries do not obviously settle fights in all groups. Territorial male butterflies, for example, compete via elaborate non-contact aerial interactions in which success is determined by relative persistence. Prior research suggests that the resolution of these contests is not clearly related to physical variables such as body size or energy reserves. However, given that the contests involve elaborate aerial manoeuvres, one long-standing suggestion is that asymmetries in flight performance, and thus flight morphology, may be important. We addressed this hypothesis via a manipulative investigation into the biophysical correlates of contest success in the speckled wood butterfly, Pararge aegeria. This species possesses the ability for significant adaptive phenotypic plasticity in relevant flight morphological parameters. We took advantage of this plasticity to rear 90 individuals of markedly varying flight morphologies, which we then pitted against each other in a semi-controlled experimental fashion. Multiple logistic and lognormal analyses provided little evidence for the relevance of morphological parameters, including relative flight musculature, wing loading and wing aspect ratio (wing length relative to area), to the outcome and/or duration of experimental contests. Instead, we found a positive effect of age upon contest success. Given that ability for high acceleration is strongly linked to variation in these morphological parameters, our findings suggest that flight performance is not a strong determinant of resource-holding potential in this notably territorial butterfly.     

Prior contest information: mechanisms underlying winner and loser effects

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

Gray tree frogs, Hyla versicolor, give lower-frequency aggressive calls in more escalated contests

As animal contests escalate, variation in the performance of aggressive signaling behaviors can give important insights into contest dynamics. In anuran amphibians, males of numerous species utilize distinctive aggressive vocalizations during disputes over calling spaces. Little is known, however, about the causes and consequences of variation in aggressive-call characteristics. We analyzed recordings of calls made during staged aggressive interactions between male gray tree frogs, Hyla versicolor, to determine how variation in a key aggressive-call characteristic, dominant frequency, was affected by increasing contest escalation. We found that dominant frequencies of aggressive calls were lower than those of advertisement calls used to attract females. Furthermore, we found that males lowered their aggressive-call frequencies with increases in escalation. Winners tended to have lower-frequency aggressive calls than losers. We conclude that aggressive calls in H. versicolor are similar to the graded aggressive calls that have been described in several other species. This gradation may allow males to balance the energetic costs of producing lower-frequency calls with the benefits of efficiently repelling rival males. Other processes related to motivation and the physiological effects of participating in contests may also be responsible for the observed variation in aggressive-call frequency with contest escalation. Our results demonstrate that detailed experimental studies of aggressive calling behavior in anurans, which to this point have rarely been performed, are feasible and generate important insights relating to general problems in animal contest behavior and animal communication.     

Symmetric fights as a measure of escalation potential in a symbiotic,territorial snapping shrimp

Summary The shrimp Alpheus armatus territorially defends the sea anemone it occupies, using as a weapon its large, specially modified snapping claw. This behavior was studied in experimental contests which were symmetric (matched individuals) with respect to sex, size, and residency. Characteristics of these contests were compared for two size-classes of male and female shrimp.There were no significant differences between these groups of shrimp in the number of bouts required to establish dominance or in the number of snaps exchanged. Large females had shorter contests than either small or large males, and losers of contests between large females were injured more frequently and more severely.This distinctiveness of large female contests could be interpreted as evidence that (i) controlling anemones is more important for large females, (ii) injuries are less important for large females, or (iii) large females lose the ability to assess one another because their contests are less frequent.If injuries are an accurate measure of the most important costs associated with fighting, then these results indicate that short contests are not necessarily the least costly, and that females can be more aggressive than males, as measured by escalation potential, in sexually selected species.     

Asymmetrical contests in defence of rock ptarmigan territories

Summary Contests involving territorial male rock ptarmigan (Lagopus mutus) occur frequently over the course of the breeding season. Fights are initiated when one male (Neighbour or Bachelor) intrudes on another's (Resident) territory. These contests involved two kinds of asymmetries: Neighbours were similar to Residents in mating status, size and experience, differing only in the gains accrued from winning a fight (payoff asymmetry) whereas Bachelors were smaller, less experienced males that had no mate or territory (fighting asymmetry, in addition to a relatively larger payoff asymmetry than in Resident x Neighbour contests). In this study, we describe and compare the contests resulting from these intrusions, and we assess how the asymmetries (fighting ability, payoffs) between opponents influenced contest intensity. Resident x Neighbour contests were brief, highly-ritualized affairs, while Resident x Bachelor interactions were long, intense battles, often ending in direct physical attack. These findings are in direct opposition to many theoretical and empirical studies on fighting asymmetries which suggest that contest intensity should be negatively correlated with the degree of fighting asymmetry between contestants. Since the potential losses associated with a contest differed between contestants and were highest for an intruding Neighbour, we conclude that payoff asymmetries were more important than fighting asymmetries in influencing a male's tendency to escalate.     

Residence duration influences the outcome of territorial conflicts in brown trout (<Emphasis Type="Italic">Salmo trutta</Emphasis>)

From the value asymmetry hypothesis, we predicted that increasing residence duration should increase perceived territory value, which should (1) increase the motivation of a replacement owner to defend the territory, (2) decrease the probability that the original owner regains the territory and (3) increase contest duration and aggression between the original and replacement owner. These predictions were tested on young brown trout. First, individual fish were allowed to establish territories for 3 days. These original owners were then moved to an adjacent territory, whereupon replacement owners were allowed to take up their territories for either 2 or 4 days. After 4 days, the original owners were returned to their old territories, now containing a new size-matched resident. The resulting conflict was observed and the initiator, duration, aggression levels and the winner of the contest were determined. Replacement trout that had been resident for 4 days initiated and won more contests against original owners than did 2-day replacements. This suggests that the motivation to defend the territory increases with residence time, which will determine the outcome of territorial contests between opponents of similar resource-holding potential (i.e. size). Moreover, contests with 4-day replacements were longer and aggression levels were higher than in contests with 2-day replacement trout. These results are consistent with the value asymmetry hypothesis, extending its validity for explaining the behaviour of territorial animals.     

Effects of ownership status,weight asymmetry,and case fit on the outcome of case contests in two populations of Agrypnia pagetana (Trichoptera : Phryganeidae) larvae

Summary The contest behavior of larvae from a dense and a sparse population of the caddisfly Agrypnia pagetana was studied in the laboratory. Fights were generally of short duration, and most fights were won by owners in both populations. Take-over contests lasted longer than non-takeover ones, indicating that owners in general used a more persistent strategy. Ownership was more important in the sparse population than in the dense population, but contest duration did not differ between the two populations. Effect of weight asymmetry was most pronounced in the dense population, and here the larger larvae usually won. However, when the intruding larvae were much larger than the owner, a low number of take-over contests were probably related to the inferior fit (low resource value) of small cases to the intruding larvae. In both populations the frequency of case takeover contests was low when the case fit to the intruder was low. We conclude that A. pagetana larvae use fighting strategies typical for contests over resources of low value relative to contest costs.     

Initiation and resolution of jumping spider contests: roles for size, proximity, and early detection of rivals

Animals are commonly expected to assess each other during contests in order to economically identify relative status. Escalated or long contests are expected to arise mainly when rivals have difficulty discriminating small differences. Results of the present study of male-male contests in Plexippus paykulli, a jumping spider (Salticidae) with acute vision, are not in accord with this widely held view. Despite the typical finding that size-advantaged rivals are more likely to win contests and that this tendency increases with size disparity, contest dynamics suggest that these tendencies are achieved in the absence of direct size assessment. In contests between different-sized spiders, maximum escalation and overall duration were predicted by the absolute size of the size-disadvantaged spider (usually the loser) rather than the size difference between the rivals. This result suggests that spiders base decisions of persistence on their own size, such that size-disadvantaged rivals usually reach their limits first, and then retreat. This interpretation is further supported by findings that maximum escalation and total duration were both positively related to size in contests between size-matched spiders. Spiders were more likely to win if they oriented and displayed first, and longer, more escalated, contests ensued if the size-disadvantaged spider was the first to orient and display. Proximity of rivals at contest outset also influenced contest dynamics, but not outcome.     

Male contests in the scarlet rosefinch (Carpodacus erythrinus) in relation to asymmetries in resource holding power and pairing status

Summary Inter-male contests in relation to asymmetries in their pairing status and resource holding power were studied in a population of the Scarlet rosefinch. Since males in this species do not defend territories before nest building, and since most males have full adult plumage, possession of territory and age are unlikely to be used as cues in the settlement of contests. Rather, two other features are used, namely size and pairing status. In this study, contests between two unpaired males were usually short displacements, whereas 56% of contests between two paired males were escalated. In the first case, the smaller male withdrew in all contests, whereas in the second case there was an equal probability of withdrawal with regard to size. There was no relationship between size difference and the probability of escalation in the contests. Most of the fights between paired and unpaired males escalated, namely in 60% of the cases where the unpaired male was smaller and in 75% of the cases where he was larger. The unpaired male withdrew in 18 out of 23 contests, suggesting that pairing status is more important than size in determining the outcome of a contest. There was no variance in size differences between contestants in short and escalated contests. This study demonstrates that there is a high competition between males for females in this population.     

Signaling effort does not predict aggressiveness in male spring field crickets

The relationship between signaling and aggression is still unclear despite several decades of research. However, there is a growing interest in studying signals that predict aggressive behavior or fighting ability. The goals of our study were threefold: we investigated the relationship between signaling effort and aggression, the relationship between body condition and aggression, and the effect of fighting experience on subsequent signaling behavior in wild-caught and laboratory-reared male spring field crickets (Gryllus veletis). We found that aggressive behavior was not related to signaling effort, body size, or body condition. For contest winners, wild-captured males were more aggressive than laboratory-reared males. Signaling effort was highly repeatable within individuals, but aggressive behavior had low repeatability. We found no evidence for a winner or loser effect on signaling; there was no change in signaling effort when we compared contest winners and losers before and after they participated in aggressive contests. Long-distance acoustic signaling and aggressive behavior appear to be independent of one another in spring field crickets, perhaps serving different functions in female attraction and male–male competition, respectively.     

Inter-sexual contests in the hermit crab <Emphasis Type="Italic">Pagurus bernhardus</Emphasis>: females fight harder but males win more encounters

Contests often occur between members of the same sex when they compete for access to mates, but inter-sexual contests may occur over access to other essential resources such as food or shelter. Despite the possibility that such contests are common, most studies focus on male fighting, and very few have analysed fights between males and females. Because males and females differ physically and physiologically, fighting ability or resource-holding potential (RHP) may also be subject to inter-sexual variation. In this study, we investigate size-controlled inter-sexual contests over the ownership of empty gastropod shells in the hermit crab Pagurus bernhardus. During these fights, there are two roles, attacker and defender, and we use a fully orthogonal experimental design to compare the performance of males and females in each role, when fighting either a male or female opponent. Although females fight more intensely, male attackers have an advantage when compared to females playing the attacker role, as they are more likely to evict the defender from its shell and thus win the resource. Further, in the defender role, male defenders are subject to shorter attacks than female defenders. The differences in agonistic performance could not be attributed to differences in perceived resource value between the sexes or to differences in body or weapon size. There are clear differences in the agonistic behaviour of males and females, and this possibility should be incorporated into models of contest behaviour. In particular, evolutionarily stable strategies may be expected to vary with sex ratios.     

Prior residency as a key determinant of social dominance in the willow tit (Parus montanus)

Summary We studied how age, body size and prior residency affected social dominance in the willow tit (Parus montanus) groups. The contribution of each variable was experimentally tested in unisexual two-bird trials, in which the birds were matched for all variables except the one studied. Large birds were dominant over smaller ones (Fig. 1). The effect of body size was more prominent in males than in females. Age had no influence on dominance. Residents became dominant more often than newcomers (Fig. 2). Adulthood or larger body size did not override the advantage of prior residency (Fig. 2). Therefore, the proximate reason for the age-dependent dominance seen in natural willow tit flocks is most likely the prior residency advantage of the adults. Factors connected with fighting ability (body size and age) seem to be less important than the time of establishment of rank, which may reflect the importance of resource value differences between residents and newcomers in this context. The advantage of residency might make it advantageous to be a member of a flock even as a subordinate, rather than being solitary.     

How do fighting ability and nest value influence usurpation contests in <Emphasis Type="Italic">Polistes</Emphasis> wasps?

Most research on animal contests has focused on the factors that influence the intensity and outcome of aggressive contests within nonsocial species, while relatively little is known about contests in social taxa. Here, we examine contests among queens of the social paper wasp, Polistes dominulus. Queens use multiple reproductive strategies, including nesting alone, usurping established colonies, and cooperatively joining other queens. We stage contests between a nesting queen and a challenger to test how resource value (RV) and resource holding potential (RHP) influence (a) who occupies the nest at the end of the contest and (b) the extent of conflict between the queen and challenger. We found that RHP, as measured by individuals’ facial patterns and body size, influenced the outcome of the contest. Challengers with high RHP were more likely to successfully usurp the nest than challengers with low RHP. Interestingly, queens with relatively high RHP were more likely to form a cooperative association with the challenger than queens with lower RHP, suggesting that queens may evict individuals that are an aggressive threat. RV influenced the intensity of conflict. There was more aggressive conflict over large nests than over small nests. Overall, social taxa have complex contest dynamics with important parallels to contests in nonsocial taxa. Studying contests in social taxa provides an important perspective on the factors that influence individual decisions about conflict versus cooperation.     

Habitat preference increases territorial defence in brown trout (Salmo trutta)

The striking ability of territory owners to repel intruders has generated a number of theoretical explanations as well as experimental studies in many animal species. However, effects of individual habitat preferences on territorial defence have rarely been studied. From the territory value hypothesis, we predicted that owners of preferred habitats should invest more resources in defence than owners of non-preferred habitats. We tested this prediction with young territorial brown trout in a two-stage experiment. First, trout were allowed to choose individually between gravel and a uniform bright substrate. As expected, they showed a significant (79%) preference for gravel. However, there was considerable variation between individuals in substrate preference, with a few fish preferring the bright substrate. Half of the tested fish were then transferred to a gravel substrate and the rest to a bright substrate, manipulating habitat type in relation to preference. Territory owners were then staged against size-matched intruders whereupon contest aggression was observed and the winner of each contest determined. Overall, owners won most of the contests. Satisfied owners won 86% and owners of less preferred territories, 74% of the contests. Furthermore, more satisfied owners attacked sooner and were more aggressive relative to the intruders. We conclude that brown trout show individual variation in habitat preference, which appears to be linked with their investment in territorial defence. These results suggest that understanding and modelling of animal contests could benefit from considering how territorial defence is influenced by individual habitat preference and specialisation. Received: 25 February 2000 / Revised: 22 May 2000 / Accepted: 25 June 2000     

When to be neighbourly: differential agonistic responses in the lizard Platysaurus broadleyi

Differential treatment of neighbours and strangers (the dear enemy phenomenon) can reduce the costly effects of territory defence. Individual recognition, and by extension the dear enemy phenomenon, was examined in the cordylid lizard Platysaurus broadleyi. I had no knowledge of familiarity between individuals and therefore classed intruders as neighbours or non-neighbours and tested for differences in agonistic response consistent with the dear enemy phenomenon. In the dear enemy phenomenon, levels of aggression are inversely related to degree of familiarity such that residents are more tolerant of immediate neighbours than distant territory holders. A manipulative field experiment in conjunction with field observations of known territory holders revealed that resident males allowed neighbours to approach more closely than non-neighbours before challenging them, residents were more aggressive towards non-neighbours than neighbours, contests between neighbours were significantly longer than between neighbours and non-neighbours and contests between residents and non-neighbours were likely to result in a win for the resident, while neighbours frequently drew contests. These results suggest a level of recognition consistent with the dear enemy phenomenon. I also manipulated front leg colour to test the hypothesis that it alone could serve as a cue for individual recognition. I found no support for this hypothesis and suggest that multiple cues operate in individual recognition. Received: 4 March 1999 / Received in revised form: 11 April 1999 / Accepted: 19 April 1999     

Sequential analysis of aggressive interactions in the stalk-eyed fly Teleopsis dalmanni

Understanding the mechanisms and determinants of conflict resolution is of great theoretical and practical importance because the outcome of contests between males over limited resources such as mates, territories, and food has profound fitness consequences. Despite the large literature on the theory of conflict resolution, relatively few empirical studies explicitly test predictions related to contest structure for these models. In sexually dimorphic species of stalk-eyed flies (Diopsidae), males engage in characteristic aggressive interactions over both females and food resources. We used sequential analysis of aggressive interactions between dyads of male stalk-eyed flies to investigate patterns of escalation, behavioral matching, and physical contact in order to distinguish between three common models of conflict resolution: the sequential assessment model, the cumulative assessment model, and the energetic war of attrition. Stalk-eyed flies were shown to engage in both low- and high-intensity behaviors during interactions with patterns of escalation and no de-escalation. Aggressive interactions did not demonstrate behavioral matching between winners and losers. Stalk-eyed flies also escalated to behaviors that included physical contact without injuries. Our results provide support for the sequential assessment model based on patterns of escalation with no de-escalation, behavioral mismatching, and behaviors which include physical contact but no injuries.