Assessment and individual recognition of opponents in the pygmy swordtails Xiphophorus nigrensis and X. multilineatus

We examined factors that determine the outcome of agonistic encounters between male pygmy swordtail fish. Xiphophorus nigrensis and X. multilineatus males formed dominance relationships based on body size in staged laboratory encounters. There was a significant negative correlation between size asymmetry and fight intensity, suggesting that males assessed size in the encounters. However, a significant proportion of the variation in fight intensity in contests that escalated to bites could not be explained by size asymmetry. Aggressive motivation may also influence the outcome of contests and could be assessed in agonistic encounters. Theory suggests that signals of aggressive intention will be evolutionarily stable if individuals can recognize opponents and encounter one another repeatedly. In addition, individual recognition is one way that dominance hierarchies can be maintained. Here we demonstrate that males from both species can recognize individuals. In addition, at least some X. nigrensis males were site-faithful in the field, suggesting males encounter the same opponents repeatedly.Communicated by G.M. Klump     

Chemical rival recognition decreases aggression levels in male Iberian wall lizards, <Emphasis Type="Italic">Podarcis hispanica</Emphasis>

In an agonistic interaction, the assessment of the probable outcome of future encounters with the same individuals may be the best way of decreasing costs of fighting, but this may only be accomplished if animals are able to recognize individual conspecifics. We staged encounters between male lizards, Podarcis hispanica, to examine whether odoriferous cues are involved in rival recognition during agonistic interactions. We experimentally manipulated the odour of intruding males, creating familiar males with their own odour or bearing odours of unfamiliar males, and unfamiliar males bearing unfamiliar odours or odours of familiar males. The results showed that when familiar males were impregnated with scents from unfamiliar males, they elicited an aggressive response by resident unmanipulated males similar to that observed for a new unfamiliar male with unfamiliar odour. This suggests that resident males were unable to recognize familiar males when their own scents were removed. In contrast, responding males were less aggressive towards familiar males impregnated with their own odour and towards unfamiliar males impregnated with scents of familiar males, suggesting that when two males have already interacted, their scents become familiar for both males, and that the detection in successive encounters of the familiar scent suffices to reduce the aggressive response of territorial males. Therefore, recognition mechanisms based on chemical cues during agonistic encounters may contribute to reducing the intensity and the costs of fighting in P. hispanica and may play an important role in the organization of their social system.     

Competitor assessment by scent matching: an experimental test

Summary If males assess potential opponents by comparing their odour with that of scent marks in the vicinity (the so called scent matching hypothesis) they should be more reluctant to fight when these odours are the same. Under these circumstances the potential opponent is thus identified as a resource holder (a male monopolising a territory or group of females) that would escalate encounters to defend its resources. We tested this prediction by comparing the fighting behaviour of male house mice, Mus domesticus, when on a scent-marked substrate that matched the odour of an opponent with their behaviour on a substrate that did not match. As predicted, the delay before the first fight was longer and the rate of subsequent fighting lower when the scents matched. This result is consistent with the interpretation that resource holders scent mark to provide opponents with a means of competitor assessment and that they thus reduce the costs of resource defence. Offprint requests to: L.M. Gosling     

No male agonistic experience effect on pre-copulatory mate choice in female earwigs

Mating with dominant males may confer considerable benefits, but also incur significant costs, hence intrasexual competitiveness is a likely target of mate choice. In addition to established modes of mate assessment, females may use cues or signals associated with agonistic experience effects to assess the relative competiveness of males. Experience effects, where the outcome of a fight increases the likelihood of a similar outcome in subsequent fights, may result from an animal’s altered state after conflict, but can also arise from strategic rival use of information perceived about this altered state. While females may similarly use this information in mate choice decisions, this potential consequence of male–male conflict has largely been neglected. Here, we investigate the effects of experience on subsequent agonistic performance in the earwig Euborellia brunneri by imposing winning or losing experiences on males and rematching them with naïve, size-matched rivals. We reveal a strong loser effect in subsequent fights, with nearly all previous “losers” losing against new rivals. In contrast, we found no equivalent winner effect, with previous “winners” exhibiting no increased likelihood of winning. We subsequently test whether the effects of male agonistic experience extend to pre-copulatory female mate choice. We show that females, when allowed to choose between naïve males and “winners” or “losers”, do not discriminate between males based on their recent agonistic experience. Therefore, while fighting history can play an important role in male–male interactions, females may not attend to this information.     

The effects of male–male contests and female eavesdropping on female mate choice and male mating success in the jumping spider, <Emphasis Type="Italic">Thiania bhamoensis</Emphasis> (Araneae: Salticidae)

In experiments that comprised of three phases (fight, choice, and mating) under “seen” and “unseen” conditions, we examined the effects of the outcomes of male–male contests and female eavesdropping on female mate choice and male mating success in the fighting spider, Thiania bhamoensis (Salticidae). The results revealed female eavesdropping on agonistic interactions. Females that had watched an aggressive interaction showed no distinctive preference for the winner over the loser, but they preferred the loser when they had not observed a fight. Winners, however, achieved a greater mating success than did losers during the mating phase. Gaining access to females was based on the insistence of the winners in courtship in terms of the number of quivers, rather than on the fighting behavior of the males. Hence, the outcome of male–male contests may not be an important determinant of a male’s mating success in T. bhamoensis. Instead, courtship display plays an important role in determining the success of male mating in this species. This study also suggests that female mate preference may not be a good indicator of eventual female mate choice and male mating success. Thus, a causal relationship between female mate preference and male mating success cannot be inferred. Joanna P. Y. Chan, Pei Rong Lau, and Ai Jie Tham contributed equally.     

Male Iberian rock lizards may reduce the costs of fighting by scent matching of the resource holders

Animals should adopt strategies to minimize the costs of intraspecific aggressive interactions. For example, individuals should be able to identify resource holders in advance and avoid fighting with them because residents are generally more likely than intruders escalate aggression. It has been suggested that scent marks function mainly to allow competitor assessment by conveying the costs of entering a scent-marked area. Individuals may identify territory owners by comparing the scent of substrate marks with the scent of any conspecific they encounter nearby, assessing whether these two scents match or not, a mechanism known as scent matching. Here, we examined the response of male Iberolacerta cyreni lizards to areas scent-marked by other males and the potential role of scent matching in agonistic interactions. We designed a laboratory experiment where we allowed a male to explore the scent-marked substrate of another male, and then we immediately staged agonistic encounters in a nearby clean neutral area with either the male that had produced the scent marks (matching treatment) or with a different non-matching individual male. The higher chemosensory exploratory rates of substrate scent marks in comparison to clean substrates suggested that males detected and spent more time exploring scent marks to obtain information on the donor male. Moreover, this information was later used to decide the fighting strategy. Intruding males delayed time until the first agonistic interaction, reduced the intensity of fights and the number of aggressive interactions, and won less interactions with males which scent matched that of scent marks (because they would be considered as the territory owners) than with other non-matching individuals. Our results show that male I. cyreni lizards use scent matching as a mechanism to assess the ownership status of other males, which could contribute to modulate intrasexual aggression, reducing costs of agonistic interactions.     

The formation and maintenance of crayfish hierarchies: behavioral and self-structuring properties

Behavioral changes in fighting and the development of dominance relationships were analyzed in groups of juvenile crayfish (Astacus astacus) using quantitative behavioral techniques. When individuals were placed into an aquarium, the number of agonistic challenges, their mean duration, and maximum intensity reached were high initially but then decreased steadily as the hierarchy developed. In all groups, linear hierarchies emerged which became increasingly stable over time. Winning influenced subsequent fighting behavior on two distinct time scales. In the short term, recent winners became progressively less likely to retreat. Second, individuals occupying dominant positions for days became increasingly likely to escalate to higher intensities early in the encounter. Both effects biased the outcome of future interactions such that winning enhanced further success and losing decreased an individual’s subsequent chances for dominance. Received: 11 November 1999 / Revised: 1 March 2000 / Accepted: 3 May 2000     

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.     

Dominant male red junglefowl (Gallus gallus) test the dominance status of other males

In sexual selection, honest signals are maintained by a variety of mechanisms. In red junglefowl (Gallus gallus), health, condition and social status affect comb size, a well-documented predictor of female choice. The comb size of subordinate male junglefowl appears to be suppressed when in the company of other males. One hypothesis for how social status could affect ornament expression in this way involves punishment of cheaters. Under this scenario, dominant males periodically challenge similar males signalling putative high status. For subordinate males, the risk of fighting a high-ranked male could make it prohibitively costly to develop ornamentation signalling dominance. We asked if dominance signals influenced the direction of aggression by dominant males. To address this issue, we conducted experiments in which 19 dominant-acting, large-combed male junglefowl were allowed to choose to fight one of two opponents. The two potential fight opponents differed in comb size, dominance behaviour, or in both traits. In 15 of 19 trials, dominant-acting males chose to fight large-combed, dominant-acting opponents rather than small-combed, subordinate-acting opponents. This is the first demonstration that aggression of dominant male birds is directed at other males based on the display of an ornament known to be attractive to females. However, males did not discriminate between fight opponents when potential opponents differed in only one of the two status indicators (large-combed males chosen in 11 of 19 trials, dominant-acting males chosen in 10 of 19 trials).     

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.     

Fighting,assessment, and frequency alteration in Blanchard's cricket frog

Summary This study examines the use of dominant frequency for assessment of fighting ability in Blanchard's cricket frog Acris crepitans blanchardi. Like most anurans, the dominant frequency of the call is negatively correlated with size. However, unlike many anurans, dominant frequency varies within individual males, providing a less reliable signal of size. Wrestling contests between male cricket frogs were differentially won by larger males. I thus tested the hypothesis that males use dominant frequency for assessment of an opponent's fighting ability. Males retreated from, or became satellites of, broadcasts of synthetic low frequency calls, simulating the calls of a large male. In contrast, males attacked broadcasts of synthetic high frequency calls, simulating the calls of a small male. The dominant frequency of the call therefore appears to contain information about size which males use to assess the fighting ability of opponents, despite the lower degree of reliability resulting from individual variation in the signal. Individual variation in dominant frequency arises partially as a consequence of active alteration of the signal in response to social competition. Males differentially lowered their dominant frequencies in response to lower frequency broadcasts, thus in response to larger opponents. Furthermore, the higher a male's dominant frequency relative to that of an opponent, the greater was the decrease in dominant frequency. These results suggest that smaller males may conditionally lower their dominant frequencies in order to sound larger to larger opponents, thereby inflating their apparent fighting ability. Two alternative hypotheses are discussed.     

Chelipeds are the real weapon: cheliped size is a more effective determinant than body size in male–male competition for mates in a hermit crab

To understand the evolution of weapons, we must understand both their functions and relative importance compared to body size in determining fighting success. Many decapod crustaceans develop disproportionately large chelipeds for their body size and use them as a weapon in agonistic interaction. There are, however, examples where weapons are merely signals of resource holding potential (RHP) and the RHP is actually determined by body size. We investigated the function and relative efficacy of body size and major cheliped size in male–male contests for females in the hermit crab Diogenes nitidimanus. Contests over females took two forms: (1) males preemptively guarded females and opponents did not fight with the guarding male. Cheliped size contributed significantly to the settlement of these contests and probably functioned as a visual signal for the opponents. (2) Guarding males engaged in physical combat with an opponent. In these cases, both body and cheliped sizes affected contest outcomes. The effect size for cheliped size was as strong, or stronger, than that for body size. These results suggest that large chelipeds have evolved as a true weapon and are effective in escalated fights for resources. Therefore they are also efficient visual signals for settling contests with only display. Our results are a rare example that clearly demonstrate that weapons are a more important determinant of fights than body size when both body and weapon size affect resource acquisition.     

Resource-holding power asymmetries,the prior residence effect,and reproductive payoffs in male northern elephant seal fights

The effect of resource-holding power (RHP) and prior residency asymmetries on fight outcome and subsequent seasonal copulatory success was analyzed for fights between marked male northern elephant seals (Mirounga angustirostris). RHP asymmetries were measured as differences in estimated mass and prior residency asymmetries were measured as differences in beach tenure prior to the fight. The principal results were: (a) Neither differences in mass nor differences in beach tenure had any effect on fight outcome as separate factors. (b) Mass and tenure differences had an interactive effect on fight outcome; fight winners were either heavier males present for shorter periods (intruders) or lighter males present for longer periods (prior residents). (c) Winners of fights copulated more often than losers after a fight throughout the breeding season; this difference was smallest for low-ranking males, larger for high-ranking males in short fights, and greatest for high-ranking males in long fights. (d) Prior resident males who won long fights obtained significantly more copulations after a fight than the males they defeated, but this was not true for intruder males who won long fights. These results suggest that male northern elephant seals will incur greater contest costs (i.e., fight for longer periods and/or against heavier males) for higher reproductive payoffs. They also imply that, at least for males in long fights, differences in prior residence represent payoff asymmetries, with higher reproductive payoffs for winning prior residents than for winning intruders.     

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.     

Differential attraction of females to dominant males: Olfactory discrimination and mating preference in the brown lemming (Lemmus trimucronatus)

Summary This study investigated differential attraction of estrous brown lemmings (Lemmus trimucronatus) to conspecific males recently exposed to each other for a 10-min agonistic encounter. In tests conducted 5 min, 1 h and 24 h after agonistic encounters, females preferred the oder of dominant males to that of defeated males when both odors were presented simultaneously in a Y-maze olfactometer. Defeat in an agonistic encounter did not reduce the propensity of male lemmings to initiate sexual behavior. In one-male, one-female tests conducted 5 min after agonistic encounters, dominant males achieved higher mount and thrust scores while defeated males obtained higher scores for attempted mounts. The sexual behavior of dominant and defeated males did not differ significantly in similar tests conducted 1 h and 24 h later. In contrast, females readily mated with dominant males and tended to avoid defeated males in two-male tethering tests conducted 5 min after agonistic encounters. In these tests, females still showed a preference for dominant males 1 h and 24 h after male agonistic encounters.     

Mate competition in the velvet swimming crab Necora puber: effects of perceived resource value on male agonistic behaviour

Male velvet swimming crabs [Necora puber (L.)] were observed to engage in long and vigorous agonistic interactions to gain possession of a sexually receptive female. The role of agonistic behaviour in male mate competition in this species was examined by investigating the effect of the perceived presence of sexually receptive females on interactions between males. Exposing male crabs to water conditioned by sexually receptive females resulted in prolonged interactions, with a greater incidence of potentially injurious behaviour than interactions between control crabs, exposed only to sea water. Maleconditioned water also resulted in more offensive behaviour, but these interactions were of shorter duration and not significantly different from sea water controls. Agonistic superiority was strongly correlated with contestant relative size when males were exposed to male-conditioned water or sea water, but not when exposed to female-conditioned water. Overall, the results conformed with the general predictions of game theoretic models in relation to the influence of resource value on agonistic behaviour. Crabs were more persistent and probably incurred greater fitness costs in the perceived presence of a sexually receptive female, when interactions may have been resolved on the basis of factors more closely related to actual fighting ability than the relative body sizes of contestants.     

Roles of male residence and relative size in the social behavior of Iberian rock lizards, Lacerta monticola

Many studies on contest competition used residency asymmetry as a discrete variable. However, the probability of winning an interaction may change as a continuous function of the value of the location where the encounter occurs. We performed a field study to examine the importance of location within a home range and relative body size to the outcomes of agonistic interactions between male lizards, Lacerta monticola. The distances to activity centers (the most used locations based on a density function of sightings) and relative size play important roles in agonistic interactions and had interacting effects in natural conditions. On the other hand, previous studies with lizards suggested that inferior competitors are able to avoid agonistic interactions in the field. Thus, we staged encounters in the laboratory to examine the behavioral responses of smaller individuals. The responses of each focal smaller male were measured in its own home cage (resident), in the cage of a larger male (intruder) and in a cage in which no male was previously present (control). The predominant behavioral tactics of smaller males were avoidance when they are the intruders and displaying when they are the residents. Submissive displays by smaller males may help reduce the costs of agonistic encounters.     

Risks and benefits of lethal male fighting in the colonial,polygynous thrips Hoplothrips karnyi (Insecta: Thysanoptera)

Summary Males of Hoplothrips karnyi (Hood) (Insecta: Thysanoptera), a colonial fungus-feeding thrips, fight each other in defense of communal egg mass sites, where they mate with females that come to oviposit. Fighting males stab each other with their enlarged, armed forelegs and hit each other with their abdomens. Escalated fights occur between large males of similar size. Fights are often lethal; males that died during observations fought more frequently than other males, were stabbed more often and more severely than other males, and were relatively large, but somewhat smaller than their opponents. Large males tend to win fights and guard egg masses, and they secure about 80% of last matings before ovipositions. Guarding males apparently assess female reproductive condition by putting their forelegs partially around females' abdomens; guarding males, but not nonguarding males, mate preferentially with females that have yet to oviposit. Non-guarding males mate with females away from egg masses, sneak matings at egg masses, and occasionally challenge guarding males. Challenges tend to follow matings by non-guarding males at egg masses. Each of four observed or inferred takeovers was followed by the death of the guarding male that lost. Male fighting strategies are discussed in terms of the consistency of lethal fighting with game theory models. Guardin males appear to pursue a classical hawk strategy of escalate until injured or victorious. This strategy may be advantageous because only large males become guarders, the mating success of guarders greatly exceeds that of non-guarders, and high population viscosity ensures that benefits from killing an opponent accrue directly to gaurders. The occurrence of challenges by large non-guarders implies that fighting ability and resource value asymmetries between males change over time; such changes may result from the energetic costs of guarding, injury to guarding males, or depletion of guarding males' supply of sperm.     

Previous experience matters in the stalk-eyed fly <Emphasis Type="Italic">Teleopsis dalmanni</Emphasis>

Previous experiences can play a significant role in determining future behaviors. Winner and loser effects, where the outcome of previous aggressive encounters influences the behavioral approach to and outcomes of future conflicts, have been documented in many taxa and illustrate this phenomenon. These effects are prevalent in species that interact frequently because modulation of these potentially costly social interactions may influence fitness. Stalk-eyed flies of the dimorphic species Teleopsis dalmanni engage in frequent fights over food resources, as well as over access to harems of females, with larger males typically prevailing when size disparities exist. However, whether and how prior experience influences fighting decisions and outcomes remains unexplored. To test for winner and loser effects in stalk-eyed flies, sexually mature flies were paired in size-mismatched dyads to establish winning and losing experiences. After their first contest, the flies were paired with size-matched individuals and allowed to interact. We determined whether an initial winning or losing experience significantly altered the outcome probabilities in the second size-matched encounter. Initial winning experience did not significantly affect the second interaction, providing no evidence for a winner effect. However, initial losers were significantly more likely to lose a subsequent interaction which provides evidence for a loser effect in stalk-eyed flies. In addition, smaller males experienced an increased probability of losing their second interaction regardless of prior winning or losing experience. This effect was not seen in large males. Our data suggest that the loser effects we observed, which were more pronounced in small males, could result from the energetic costs of fighting that they were less able to absorb than large males.     

Territorial counter-singing in male sac-winged bats (<Emphasis Type="Italic">Saccopteryx bilineata</Emphasis>): low-frequency songs trigger a stronger response

In many animal species, individuals compete for resources but avoid escalated conflicts by threat displays, i.e. a mutual signalling behaviour that enables the opponents to predict the outcome of the conflict without the necessity of actual fighting. For example, territory holders may use acoustic signals to communicate not only their own identity and the borders of their territory but also their competitive quality, fighting ability and motivation. Here, we show that male sac-winged bats, Saccopteryx bilineata, adjust their vocal territorial displays according to the fundamental frequency of territorial songs of their opponents. In playback experiments with territorial males, low-frequency stimuli elicited a higher territorial song rate and length than high-frequency stimuli. Male S. bilineata that sing more often and with lower fundamental frequencies have been shown to sire more offspring than their competitors. Fundamental frequency of territorial songs, hence, may reveal male quality and, consequently, the resulting threat posed to competing males. We argue that this is reflected in the increased response of competitors to low-frequency territorial songs shown here. Such competitive signalling behaviour has been shown in a few mammal species like red deer and baboons but, thus far, not in bats. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.