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.     

Do Midas cichlids win through prowess or daring? It depends

Summary Individual Midas cichlids (Cichlasoma citrinellum) show persistent differences in aggressive behavior toward dummy fish. The starting question was whether the level of such behavior can be used to predict the winner of a fight. In the first (long-term) experiment two fish were matched for size, color, and sex. After 24 h the opaque barrier separating the 2 fish was removed; they immediately behaved aggressively. Scores for aggression toward dummies did not predict the winners, nor did taking the initiative in escalation. But weight did foretell the winners, who averaged only 2% heavier than their opponents. The conventional display phase of such fights was brief (20% of total duration), and escalation was rapid. Both winners and losers sustained damage, but losers accumulated damage faster than winners. In the subsequent (short-term) experiment the fish were separated only 1–2 h. Now aggression scores predicted winners, and winners were the fish who escalated. Weight of fish had no effect. The conventional phase was relatively much longer, about half the length of the fight. Losers accumulated damage at the same rate as the losers in the long-term experiment, but the fights were shorter; winners suffered little damage. The fish had difficulty assessing one another. Fighting prowess was remarkably uniform when weight was factored out. Daring to escalate, in contrast, varied among individuals and correlated with aggression scores. Prowess (=weight) determined the outcome in the long-term experiment, which may find its parallel in nature in intraterritorial disputes. Daring to escalate determined the winner in the short-term experiment; this may be comparable to establishing a territory in nature.Prowess probably results from strong directional selection because it has low costs and high benefits. In contrast, daring is subject to bi-directional selection because both costs and benefits are high. Resource holding potential is conventionally viewed as deriving from prowess of self and opponent and value of resource; to that one must add the individual's inherent aggressiveness.     

Defeated chameleons darken dynamically during dyadic disputes to decrease danger from dominants

Research on intraspecific aggression has typically focused on dominant individuals, but a better understanding of the consequences and mechanisms of agonistic encounters requires a balanced perspective that includes knowledge of subordinate animal behaviors. In contrast to signals of fighting ability, signals of submission are an understudied component of agonistic communication that could provide important insights into the dynamics, function, and evolution of intraspecific competition. Here, I use a series of staged agonistic trials between adult male veiled chameleons Chamaeleo calyptratus to test the hypothesis that rapid skin darkening serves as a submissive signal to resolve agonistic activity. Concordant with this hypothesis, I found that losing chameleons darkened over the course of aggressive trials while winners brightened, and the likelihood of darkening increased when individuals were attacked more aggressively. Additionally, I found that the degree of brightness change exhibited by individual chameleons was tied to both overall and net aggression experienced during a trial, with chameleons who received high levels of aggression relative to their own aggression levels darkening to a greater extent than individuals receiving relatively less aggression. Lastly, I found that aggression increased for losers and winners prior to the onset of darkening by the eventual loser but that both chameleons reduced aggression after the losing chameleon began to darken. Based on the theoretical prediction that signals of submission should be favored when retreat options are restricted, I suggest that limited escapability imposed by chameleon morphology, physiology, and ecology favored the evolution of a pigment-based signal of submission in this group.     

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.     

Soft song is a reliable signal of aggressive intent in song sparrows

Animals frequently use signals to modulate aggressive interactions. Establishing that a signal is aggressive or threatening requires demonstrating that it is more commonly used in agonistic contexts, that it predicts subsequent aggressive behaviors by the sender, and that receivers respond differently to this signal. Like many birds, song sparrows (Melospiza melodia) produce a low-amplitude “soft song” vocalization that has been hypothesized to be an aggressive signal. Soft song meets the first two criteria, but previous research has failed to demonstrate that soft song provokes aggression or that receivers even perceive soft song differently from normal loud song. We used a playback experiment with taxidermic mount presentation to test whether territorial male song sparrows respond differently to loud and soft song playbacks. Subjects reacted more strongly to the soft song playback by approaching the mount more closely, increasing wing wave displays, and increasing the proportion of their own songs that were soft songs, with further trends toward increasing the number of flights and attacks. These results confirm that soft song is a conventional signal of aggression in song sparrows and that increased receiver retaliation maintains its reliability.     

Immune function, dominance and mating success in drumming male wolf spiders Hygrolycosa rubrofasciata

In this study, we provide a piece of experimental evidence that immune function is related to dominance and mating success in wild caught male wolf spiders, Hygrolycosa rubrofasciata. In the mating season, H. rubrofasciata males are actively searching for receptive females, and while searching males often engage in agonistic behavior (i.e., agonistic drumming signals, chases, and fights) with each other. The present results demonstrate that dominant males had higher lytic activities in their hemolymph than subordinates. Lytic activity estimates the concentration of antimicrobial peptides with lysozyme-like activity in hemolymph, which have been shown to play an important role in defense against bacteria, viruses, and fungi. Dominants also had higher courtship drumming rates than subordinates. Moreover, winners in mating competition had higher lytic activities than losers, but this was measured nonindependently of dominance status. Among males with mating failure, there was a moderate negative correlation between encapsulation rate and courtship drumming rate, suggesting that low quality males might not be able to bear the immunological costs of courtship behavior. These results suggest that females might gain immunological benefits by preferring vigorous males.     

Escalating fights for rank-order position between male swordtails (Xiphophorus helleri): Effects of prior rank-order experience and information transfer

Summary A previous hypothesis which we originally based on standardized mirror tests was examined. It stated that subordinate males from socially stabilized groups, when confronted with a strange male of similar fighting ability, tend to take the offensive; in contrast, dominant males tend to wait and see and to increase their aggressiveness only as far as necessary to counter the aggressive actions of the opponent. 30 highly escalating fights between well matched opponents, either with different or with the same prior rank-order experience, were recorded on video tape and then analysed in detail. Rank-order experience affected neither the escalation level nor the outcome of the fights. However, in the first nonescalated stage of the fights all measures of unilateral aggressive actions were higher in the omega than in the alpha males. This is contrary to expectations based on the theory of social conditioning; nor can it yet be explained by game-theory models. We then discuss why it should pay subordinates to be more offensive and to show higher levels of unilateral aggression than dominants. Some current aspects of game theory are supported by our data: (1) Threat display is a poor predictor of attack and fleeing, and simply informs the opponent that the threatening individual will probably just stay and avoid further escalations. (2) Relative fighting ability is demonstrated during highly-escalated fighting by delivering Fin Grips, the most effective and costly attack pattern. The high energetic costs of Fin Grips can be compensated by reducing the rate of Biting. (3) Probably, the combatants did not transfer information about their intentions, i.e. they concealed their intentions as to what they were going to do next. Contrary to the predictions of game theory, winners and losers showed some differences in Pecking early in the nonescalated stage of the fights, the prospective losers exhibiting more Pecking than the winners.This study was supported by grants from the Deutsche Forschungsgemeinschaft     

Females prefer males with superior fighting abilities but avoid sexually harassing winners when eavesdropping on male fights

Selection imposed by male competition (intrasexual selection) and female choice (intersexual selection) can be con- or discordant. Specifically, females may or may not prefer mating with dominant males, and direct costs of interacting with dominant (and possibly more harassing) males have been suggested to explain avoidance of dominant males. Here, we exemplify that inter- and intrasexual selection may normally act in the same direction, but can be temporarily conflicting when social information becomes available. Using video playback techniques, we presented females of the Mexican livebearing fish Poecilia mexicana with two size-matched males and established association preferences. Half of the females could then observe the same two males fight and establish dominance, while control females saw both males side by side, but physically separated, and female preferences were subsequently re-evaluated. Females in the control group showed a significant preference for future winners in the subsequent testing, confirming an innate or acquired preference for male traits that are indicative of physical superiority, even when body size as a choice criterion is excluded. When allowed to eavesdrop on male fights, however, females did not show a preference for observed winners and even decreased time spent with them relative to the control treatment in which no fight was shown. A subsequent experiment found contest winners to show elevated levels of sexual behavior, so we argue that the temporary offset of the intrinsic female preference for dominant males after having observed a fight is indeed driven by direct costs females expect from more harassing contest winners.     

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.     

Past ownership makes crayfish more aggressive

There is plenty of evidence that resource value is one of the most important non-strategic variables in animal fighting behavior. Here, we tested whether the past ownership of a shelter might modify the agonistic behavior of the crayfish Austropotamobius pallipes, eventually increasing its probability to win when it reencounters a previously met conspecific away from that resource. The agonistic behavior of familiar pairs composed of size-matched males was observed for an hour; after that, the two contestants had been kept in isolation for 2 days, either in the presence or in the absence of a shelter. Specifically, in the isolation phase, a shelter was offered to (1) both crayfish, (2) no crayfish, (3) the dominant crayfish only, and (4) the subordinate crayfish only. The following combat was conducted in the absence of any refuge. The crayfish that previously owned a shelter showed a higher aggressive motivation to fight than the individuals kept without a shelter. Particularly, in the pairs (4), subordinate crayfish were even more aggressive than dominants but were never able to invert hierarchies. Taken together, our results confirm the role played by shelters as determinants of agonism and also show, for the first time, how the behavior of crayfish and their internal state may be affected by their past ownership of a resource.     

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.     

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.     

The role of size and aggression in intrasexual male competition in a social lizard species, Egernia whitii

Competition between males is a key component of the agonistic intrasexual interactions that influence resource acquisition, social system dynamics, and ultimately reproductive success. Sexual selection theory predicts that traits that enhance success in intrasexual competition (particularly male–male competition) should be favored. In vertebrates, this often includes body size and aggression, with larger and/or more aggressive males outcompeting smaller or less aggressive conspecifics. The majority of studies consider aggression as a flexible trait which responds to local social or environmental conditions. However, aggression frequently shows considerable within-individual consistency (i.e., individuals have identifiable aggressive behavioral types). Little is known about how such consistency in aggression may influence competition outcomes. We integrated a detailed field study with a laboratory experiment to examine how a male’s aggressive phenotype and his size influence competitive interactions in Egernia whitii, a social lizard species which exhibits strong competition over resources (limited permanent shelter sites and basking sites). Individual aggression and size did not predict competition outcome in the laboratory nor did they predict home range size, overlap, or reproductive success in the field. However, winners of laboratory trial contests maintained consistent aggressive phenotypes while consistency in aggression was lost in losers. We suggest that aggression may be important in other functional contexts, such as parental care, and that alternative traits, such as fighting experience, may be important in determining competition outcome in this species.     

A new heuristic for capturing the complexity of multimodal signals

Many animal signals are inherently multimodal, engaging more than one of the receiver’s sensory systems simultaneously, and it is the interaction between the two modalities that determines the signal’s function (s) and efficacy. It is hence necessary to quantify the effect of each modality relative to the other in order to fully understand animal communication. We have developed a new heuristic to aid in the identification and interpretation of the many distinct ways in which signals in multiple sensory modalities interact. Our approach represents natural variation in signal production for each modality and uses these to generate three-dimensional receiver response surface plots that map the relationships among the signal components and receiver behavior. We accommodate the extant hypotheses for the interactions between modalities, each of which makes a clear prediction about the shape of the response surface, and extend previous theory by considering new phenomena.     

Social control of deception among status signalling house sparrows Passer domesticus

Summary Motivation to fight and willingness to risk the use of dangerous weapons, two factors which can be decisive in predicting the winners of asymmetric animal contests, are readily bluffed in display. When contests are decided by motivation to fight or daring, more reliable information about contest asymmetries may be gained by watching actual fights than by watching displays. By presenting mounts to territorial red-winged blackbirds and scoring how aggressively a territory holder attacks the mount, then recording any subsequent intrusions over established boundaries, I showed that neighbors are likely to intrude upon focal males that fail to attack the mount vigorously. This result suggests that red-winged blackbirds watch contests to assess the RHP of neighbors.     

Adaptive strategies of territory formation

How do territorial animals gain ownership of an area? Early modelling has considered the evolution of fighting when the winner can claim the right to the resource. Recently, alternative hypotheses have been offered where repeated interactions lead to division of space through 'nagging' instead of one decisive fight. However, these models assume that animals avoid areas in which they have taken part in aggressive interactions, but do not consider whether avoidance itself is adaptive. We aim to bridge this gap between mechanistic and adaptive explanations, by presenting a game-theory model where individuals choose whether to return to an area after a fight with a specific outcome (win, loss, draw). We show that avoidance of areas where fights have occurred can be adaptive, but only if benefits of access to the area are low compared to costs of fighting. Otherwise, one individual (typically the winner) responds by returning to the area, and the other (loser) avoids it. In such cases, space is gained by winning fights. We also consider the role of conventions. If responses to fights were purely conventional, paradoxical strategies where losers of fights gain ownership would be equally logical as common-sense ones where winners claim ownership. Paradoxical solutions can be stable but only when there is little difference in fighting ability between the competitors, when individuals adhere very strictly to a behavioural rule without much random variation, and when the population in its ancient state used a paradoxical strategy.     

Chorus model of the synchronizing bushcricket species Mecopoda elongata

Males of the Malaysian bushcricket species Mecopoda elongata synchronize or alternate their cyclically occurring song elements (chirps) in a duet. The acoustic interaction of males interacting in a duet was successfully simulated by means of mutually coupled song oscillators, which respond to a disturbance by a phase shift which is known from the phase response curves (PRCs) of real males. However, little is known about the acoustic interaction of males in a complex chorus situation. Therefore, the aim of the current study was to extend the duet model to a chorus taking into account an inhomogeneous spacing of agents and a natural variability of oscillator properties. This chorus model was used to study oscillator coupling in a chorus consisting of 15 agents. Since such a computer model allows one to simulate chorus manipulations that far exceed the possibilities of behaviour experiments, the following scenarios were simulated: modification of chorus density, sensory bias during sound production, selective attention to only a subset of neighbors and males joining or leaving a chorus. Simulation results allow one to draw conclusions about the chorusing behavior of males in a real chorus and about signaler and receiver aspects influencing chorusing formation and mate choice.     

How a songbird with a continuous singing style modulates its song when territorially challenged

In songbirds of the temperate zone, often only males sing and their songs serve to attract females and to deter territorial rivals. In many species, males vary certain aspects of their singing behavior when engaged in territorial interactions. Such variation may be an honest signal of the traits of the signaler, such as fighting strength, condition, or aggressive motivation, and may be used by receivers in decisions on whether to retreat or to escalate a fight. This has been studied intensively in species that sing discontinuously, in which songs are alternating with silent pauses. We studied contextual variation in the song of skylarks (Alauda arvensis), a songbird with a large vocal repertoire and a continuous and versatile singing style. We exposed subjects to simulated territorial intrusions by broadcasting conspecific song and recorded their vocal responses. We found that males sing differently if they are singing spontaneously with no other conspecific around than if they are territorially challenged. In this last case, males produced lower-frequency syllables. Furthermore, they increased the sound density of their song: they increased the proportion of sound within song. They seem to do so by singing different elements of their repertoire when singing reactively. Furthermore, they increased the consistency of mean peak frequency: they repeated syllable types with less variability when singing reactively. Such contextual variation suggests that skylarks might use low frequencies, sound density, and song consistency to indicate their competitive potential, and thus, those song features might be important for mutual assessment of competitive abilities.     

Weapon size versus body size as a predictor of winning in fights between shore crabs, Carcinus maenas (L.)

Relative body size (carapace width) and weapon size (chela length) were used as indicators of resource holding potential (RHP) in the agonistic behaviour of male shore crabs, Carcinus maenas (L.). Weapon size was found to be a more reliable predictor of the outcome of pairwise fights than body size. Crabs with longer chelae than their opponents were more likely to win fights than crabs with relatively larger bodies. Body size had less influence on the outcome of fights. Relative body and weapon size did not influence initiation of contests but did affect the likelihood of winning; however, this was significant only for weapon size. Winning crabs had heavier claws with greater surface area than losing crabs. There was no relationship between relative size and fight duration. The frequency of cheliped display increased with chela length and win- ners performed significantly more displays than losers. Received: 5 February 1997 / Accepted after revision: 20 May 1997     

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.