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.     

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     

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.     

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.     

Payoff asymmetries in contests among male brown-headed cowbirds

Summary Morphometric analyses and behavioral observations of a marked population of brood parasitic brown-headed cowbirds (Molothrus ater) revealed several male attributes correlated with dominance. Males that won larger percentages of the contests in which they were involved tended to be older and larger than other males. Dominant males arrived earlier on the breeding grounds and became engaged in more contests than subordinate males. If a female was present during a contest among males, the male which she was accompanying (her consort, and mate in most cases) was more likely to win the contest than were any of his opponents. Multiple regression analyses were performed to determine any significant partial effects of these attributes on male dominance. Age and involvement in more contests exerted significant partial effects in some analyses, but only the presence of a female consort did so consistently. I suggest that the female is a resource of greater value to the male with whom she is consorting than to other males. Her value is greater in terms of present and future benefits because he has already expended some or all of the effort necessary to establish a pair bond. He will have to expend less future effort to remain her mate than would another male to become her mate. Her presence during contests induces him to persist (or escalate) and to dominate his male opponents.     

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.     

Exaggerated male eye span influences contest outcome in stalk-eyed flies (Diopsidae)

Evolution of male weapons or status signals has been hypothesized to precede evolution of female mating preferences for those traits. We used staged male fights among three species of Malaysian stalk-eyed flies (Diptera: Diopsidae) to determine if elongated eye span, which is preferred by females in two sexually dimorphic species, influences contest outcome. Extreme sexual dimorphism, with large males possessing longer eye span than females, is shared by Cyrtodiopsis whitei and C. dalmanni. In contrast, C. quinqueguttata exhibits a more ancestral condition – short, sexually monomorphic eye stalks. Videotape analysis of 20-min paired contests revealed that males with larger eye span and body size won more fights in the dimorphic, but not monomorphic, species. To determine if males from the dimorphic species use eye span directly to resolve contests, we competed male C. dalmanni from lines that had undergone artificial selection for 30 generations to increase or decrease eye span. We found that eye span, independently of body size, determines contest outcome in selected-line males. Furthermore, in both dimorphic species, the average encounter duration declined as the eye span difference between contestants increased, as expected if males use eye span to assess opponent size. The number of encounters also increased with age in dimorphic, but not monomorphic, species. Selected-line males did not differ from outbred males in either fight duration or number of encounters. We conclude that exaggerated male eye stalks evolved to influence both competitive interactions and female mating preferences in these spectacular flies. Received: 20 July 1998 / Received in revised form: 2 February 1999 / Accepted: 12 March 1999     

Twilight fighting in the evening brown butterfly,<Emphasis Type="Italic"> Melanitis leda</Emphasis> (L.) (Nymphalidae): age and residency effects

Butterflies are conspicuous among animals that fight for mating opportunities because it is not clear how contest costs could accrue. Nevertheless, the bulk of research in this group suggests that contests are settled on the basis of asymmetries in fighting ability (the superior competitor hypothesis). There is also a consistent effect on contest escalation due to apparent 'confusion' over residency, which could result from resident butterflies playing different tactics than non-residents, or from a causal link between residency and fighting ability. I addressed these hypotheses by studying contest behavior in the crepuscular species Melanitis leda (L.) (Nymphalidae) over a 5-month period in tropical Australia. Males competed via conspicuous two-stage maneuvers of a form unique among butterflies. Prior residents won 77% of all contests. Non-resident males arrived and perched in occupied sites until challenged by incumbent residents, and the aggressiveness of these interlopers increased as a function of the time before they were detected. Contest winners also tended to be younger than losers, and contest escalation was negatively related to both the age of the losing male and the magnitude of the between-combatant age asymmetry. These results are consistent with the superior competitor hypothesis based upon age as a determinant (or correlate) of fighting ability, but also suggest residency has a unique influence on contest behavior. Further research is clearly required to unravel the effects of age, residency and other potential biophysical determinants of fighting ability in this species.     

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.     

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.     

Intraspecific aggression in the colour morphs of the anemone Phymactis clematis from Chile

The anemone Phymactis clematis exhibits intraspecific agression. An investigation into dominance relationships amongst the 3 colour morphs collected from Vna del Mar, Chile, during June, 1980, revealed them to be equally aggressive. The outcomes of experimental contests appeared to reside solely with size-dependent differences in the threshold for the release of agression. Contests were asymmetric and larger individuals displayed aggression earlier against smaller opponents, and won encounters. An analysis of times for the onset of aggression in contests showed that they were highly variable and were not a good indicator of the absolute aggressive ability of individuals as denoted by dominance rankings. It provided, however, evidence indicating a lack of assessment of opponent size during initial contact. This is discussed in relation to the evolution of aggression in anemones and to the concept of evolutionarily stable strategies in contest behaviour. It is concluded that future work on the apparent size-dependent threshold will most instructively be conducted at the neurophysiological level.     

The production of sexual and asexual larvae within single broods of the scleractinian coral, Pocillopora damicornis

The genotypic compositions of two populations of the brooding coral, Pocillopora damicornis, were studied by exhaustive sampling of adult colonies on a fine scale in southern Taiwan. In addition, 100 larvae were randomly selected among more than 1,000 brooded larvae collected from a single broodparent within each population. Using 7 polymorphic microsatellite markers, both populations were found to be highly clonal, and 7 clonal lineages were characterized. One clonal lineage (C1) dominated both study areas and comprises 54.9% of all colonies sampled, while any of the other 6 clonal lineages represented no more than 5%. Among the 100 larvae randomly selected for genotyping from each broodparent, the extent of clonal reproduction was high, and only 29 and 6 larvae, respectively, were found to be genotypically different from their broodparent. Among the 35 genotypically distinct larvae, 33 were thought to be derived from sexual reproduction, and 2 were assumed to be clonal propagules that had undergone somatic mutations. Two genotypically identical larvae were also found in one of the 2 sexually derived larva arrays, indicating the possible existence of polyembryony. The high proportions of clonality in both adult colonies and brooded larvae suggest that asexually produced larvae might significantly contribute to the recruitment of local populations. The dense clonal population of P. damicornis in the study area favors the quick recolonization view of clonal propagules after disturbances.     

The effect of size and cheliped autotomy on sexual competition between males of the mud crab <Emphasis Type="Italic">Cyrtograpsus angulatus</Emphasis> Dana

Size advantage in male–male competition over mates, combined with male preference over large females, is a common feature that can drive to size assortative mating and, eventually, sexual selection. In crabs, appendage autotomy can affect assortative mating and opportunity for sexual selection by affecting size advantage in mating contests. In this work, we evaluate the effect of size and appendage autotomy in generating assortative mating in the mud crab Cyrtograpsus angulatus. Field observations of guarding pairs in two different populations show a positive correlation between carapace width of males and females in both the populations. In one of the populations, incidence of appendage autotomy was low and the variability in the size of reproductive males was lower than the variability in the size of randomly collected males (i.e. only larger males were successful in getting a female), whereas there was no differences in the other population (i.e. most male sizes were successful) where the incidence of appendage autotomy was very high, indicating that the importance of size is higher when the incidence of autotomy is low. In this context, experiments (in both populations) show that, in contests for a female, larger males outcompete smaller ones only when they had intact appendages. When males had missing chelipeds, winning or loosing against smaller males was random. This may lead to a decrease in the importance of male size in populations with high incidence of cheliped autotomy, affecting assortative mating and opportunity for selection and, thus, affecting selective pressures.     

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.     

The outcome of in-vitro contests between larvae of the endoparasitoid<Emphasis Type="Italic"> Venturia canescens</Emphasis> depends on both their relative and absolute ages

In a laboratory colony of the asexual solitary endoparasitoid Venturia canescens Grav. (Hymenoptera: Ichneumonidae), two genetically and phenotypically distinct lines (RP and RM) appear to coexist sympatrically, with the reproductive success of the RM-line being greater under con-specific superparasitism but lower under self superparasitism. To investigate the basis of the RM-lines advantage under con-specific superparasitism, we examined the outcome of inter-larval physical combat under in-vitro conditions. The results showed that the outcome depended on both the relative and absolute ages of the contestants, and that the competitive abilities of the two lines were not symmetric. In contests involving a larva and an egg, the unhatched wasp lost. In contests involving two larvae, at least one of which was newly hatched, the RP-larva tended to lose. Finally, if both larvae were at least 8–10 h post-hatching when the contest occurred, then the larger larva tended to lose, with the probability increasing with the difference in length between the two larvae. Thus, the higher reproductive success of the RM-line under competing superparasitism with the RP-line is due to a physiological difference between the newly hatched larvae of the two lines, which results in an advantage to the RM-larva independent of the order or time interval between ovipositions.Communicated by P. Heeb     

Perceived predation risk and mate defense jointly alter the outcome of territorial fights

Virtually all animal conflicts occur over access to mates or resources that affect survival, the two key components of fitness. In this paper, we report that predation risk and mate defense jointly affect the outcomes of contests between male sand crabs (Scopimera globosa) for burrows in which crabs mate and take shelter from predators. We observed the contests under three different conditions: (1) the natural condition of low predation risk and without the presence of a female; (2) the first experiment in which we imposed upon only intruding males the perception of predation risk—by digging them from their burrows, capturing and handling them, and placing them into other males’ burrows—to increase the value of the burrows for the intruders as shelter, and (3) the second experiment in which we repeated this treatment but increased the resource value of the burrow to the resident by placing a female in his burrow. The difference in body size between contestants was the main determinant of victory in all analyses. However, perceived predation risk also partly affected the outcomes of the fights: The motivated intruders were likely to win even when they were a little smaller than the residents. In addition, defense of a female had a significant effect on the outcomes of fights: The motivated residents won more fights than the motivated intruders, indicating that these two treatments caused asymmetric increases of the resource value. This is the first report of two external factors simultaneously raising resource value, affecting motivation of contestants, and altering the outcome of fights.     

The value of patches of intertidal seagrass to prawns depends on their proximity to mangroves

Penaeid prawns were sampled with a small seine net to test whether catches of postlarvae and juveniles in seagrass were affected by the distance of the seagrass (mainly Zostera capricorni) from mangroves and the density of the seagrass in a subtropical marine embayment. Sampling was replicated on the western and eastern sides of Moreton Bay, Queensland, Australia. Information on catches was combined with broad-scale spatial information on the distribution of habitats to estimate the contribution of four different categories of habitat (proximal dense seagrass, distal dense seagrass, proximal sparse seagrass, distal sparse seagrass) to the overall population of small prawns in these regions of Moreton Bay. The abundance of Penaeus plebejus and Metapenaeus bennettae was significantly and consistently greater in dense seagrass proximal to mangroves than in other types of habitat. Additionally, sparse seagrass close to mangroves supported more of these species than dense seagrass farther away, indicating that the role of spatial arrangement of habitats was more important than the effects of structural complexity alone. In contrast, the abundance of P. esculentus tended to be greatest in sparse seagrass distal from mangroves compared with the other habitats. The scaling up of the results from different seagrass types suggests that proximal seagrass beds on both sides of Moreton Bay provide by far the greatest contribution of juvenile M. bennettae and P. plebejus to the overall populations in the Bay.Communicated by M.S. Johnson, Crawley     

Do the elongated eye stalks of Diopsid flies facilitate rival assessment?

The elongated eye span of male Diopsid flies is a sexually selected character that scales positively with body size. Previously, the duration of agonistic contests was found to increase as rival body size and eye span disparities decreased. Hence, along with its role in mate choice, eye span seems to facilitate mutual assessment of rival size. However, such results are also expected in the absence of rival assessment, when each individual persists according to its own size-dependent internal threshold. Here, we reanalyze these contests to distinguish between these two hypotheses using two measures of size: body length and eye span. Mutual assessment predicts that contest duration should increase with loser size and decrease with winner size. In contrast, our results were more consistent with self-assessment: We found a positive relationship between loser size and contest duration, whereas winner size did not affect contest duration. Thus, flies did not appear to assess the size of their rivals, indicating that the mutual assessment function of eye span elongation may be less important than previously suspected. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Inbreeding and competitive ability in the common shrew (<Emphasis Type="Italic">Sorex araneus</Emphasis>)

Common shrews (Sorex araneus) maintain a foraging territory for most of their immature life. Possessing a high-quality territory is vital for overwinter survival in the harsh boreal climate, and hence, competitive ability in territorial disputes is expected to be an important component of individual fitness. To test possible association between individual inbreeding and fitness, we used neutral arena trials to assess the competitive performance of young common shrews. The experiment involved pairs of individuals originating from small island populations, where breeding must often occur between related individuals, and from large outbred mainland populations. The percentage of neutral arena tests that an individual won was highly significantly explained by internal relatedness, a surrogate measure of individual inbreeding, measured using ten microsatellite markers. Body size, sex, learning, and population type (mainland vs island) made no significant contributions. Even a low level of individual inbreeding may lead to significant adverse consequences in multiple territorial contests, which may represent a significant cause of inbreeding depression in many wild vertebrate populations.     

Direct male-male competition can facilitate invasion of new colour types in Lake Victoria cichlids

The possibility that disruptive sexual selection alone can cause sympatric speciation is currently a subject of much debate. The initial difficulty for new and rare ornament phenotypes to invade a population, and the stabilisation of the resulting polymorphism in trait and preference make this hypothesis problematic. Recent theoretical work indicates that the invasion is facilitated if males with the new phenotype have an initial advantage in male-male competition. We studied a pair of sympatric incipient species of cichlids from Lake Victoria, in which the red (Pundamilia nyererei) and blue males (P. pundamilia) vigorously defend territories. Other studies suggested that red phenotypes may have repeatedly invaded blue populations in independent episodes of speciation. We hypothesised that red coloration confers an advantage in male-male competition, assisting red phenotypes to invade. To test this hypothesis, we staged contests between red and blue males from a population where the phenotypes are interbreeding morphs or incipient species. We staged contests under both white and green light condition. Green light effectively masks the difference between red and blue coloration. Red males dominated blue males under white light, but their competitive advantage was significantly diminished under green light. Contests were shorter when colour differences were visible. Experience of blue males with red males did not affect the outcome of a contest. The advantage of red over blue in combats may assist the red phenotype to invade blue populations. The apparently stable co-existence of red and blue incipient species in many populations of Lake Victoria cichlids is discussed.Communicated by C. St. Mary