Resource Holding Potential and the Outcome of Aggressive Interactions between Paired Male <Emphasis Type="Italic">Aegus chelifer chelifer</Emphasis> (Coleoptera: Lucanidae) Stag Beetles

Interactions between male stag beetles usually involve aggressive behavior using their long mandibles as weapons to compete with rival males over females. Considerable variation exists within populations in male body size, and may affect their behavior and the outcome of male-male contests. We investigated the aggressive interactions between male Aegus chelifer chelifer, a small tropical stag beetle species. Morphological traits in relation to aggressiveness and the outcome of fights were examined in laboratory-reared beetles. The fight-engagement ratios of major and minor morph males were not significantly different and analyses revealed that the size of body parts had more effect on the fighting success than the weapon part (mandibles). The probability of winning a contest was higher in males with a larger head width (HW), and so HW was considered as the resource holding potential (RHP). No effects of the trait size on the initiation of fights or aggressive intensity was found. Relationships between the fight duration and RHP were not significantly consistent with any assessment strategies, but were close to the mutual assessment model.     

Male contest investment changes with male body size but not female quality in the spider Nephila clavipes

Animals use rules to adjust their level of investment in a contest. We evaluate male strategies during contests over females in the golden orb-web spider Nephila clavipes. We tested whether male behaviour changes with female value, and found that contests were similar in intensity and outcome whether the female was a juvenile or adult, virgin or non-virgin, or whether one male had invested sperm in the female. We found evidence that males use a self-assessment strategy when deciding to withdraw from a contest. Loser body size and contestant size difference were correlated with a higher frequency of contest escalation, and fights involving two large males were more likely to escalate than a fight in which one male was small. A multiple regression showed that loser body size had a stronger effect on contest escalation than contestant body size difference. More importantly, the size of the winning male had no effect on contest escalation, a key prediction of a self-assessment strategy. In N. clavipes, body size is the primary factor that determines the outcome of male contests, and males do not appear to assess their opponent or the quality of the resource when deciding to withdraw from the fight.     

Aggression and RHP in the Northern Swordtail Fish, Xiphophorus cortezi: The Relationship Between Size and Contest Dynamics in Male–Male Competition

Current theory predicts that contest outcome, as well as decisions on whether to initiate a contest, escalate during a contest or retreat are decided by asymmetries in resource holding potential (RHP) and/or expected payoffs between contestants. In this investigation, dyadic contests were staged between male swordtail fish (Xiphophorus cortezi) where individuals were paired based on cumulative fight records and were ranked at the end of the trials in order to approximate RHP. Size was the only asymmetry that I did not attempt to control for and as a result, I was able to determine the relationships between size, contest initiation, escalation and outcome. Individuals changed their contest initiation strategy based on their size relative to that of their opponents, and contrary to predictions, the smaller of the two males in each contest was more likely to initiate the conflict than was the larger male. However, the larger of the two males was more likely to win and standard length proved to be a moderate predictor of an individual's final rank. Regardless of size, initiators fared poorly, winning only 31% of the contests. In instances where the smaller males won the contests, they were no more likely to initiate the encounter than was the larger male. However, when small males did win, fights lasted longer, suggesting that in some cases smaller males may be able to outlast their opponents.     

Fine‐ and Broad‐Scale Approaches to Understanding the Evolution of Aggression in Crickets

Male field crickets frequently engage in agonistic contests to establish dominance in social interactions and gain access to mate attraction territories. Crickets (Orthoptera: Gryllidae) are often used as a model taxon to study aggression, but limited documentation of aggression in some cricket species hinders our understanding of its evolutionary costs and benefits. Our study investigated cricket aggression at two scales: the within‐species scale for two cricket species, Gryllus assimilis and G. veletis, whose aggression had not been adequately documented and the among‐species scale to detect evolutionary patterns in species’ levels of aggression. In both G. veletis and G. assimilis, winners spent more time being aggressive than losers, but they were not larger or heavier. Collectively, our results reveal that G. veletis males are more aggressive than G. assimilis. Male G. veletis had higher aggression scores that male G. assimilis. The majority of G. veletis contests escalated to grappling (a highly aggressive behavior), while less than one quarter of G. assimilis contests escalated to grappling. Further, G. veletis males transitioned between two of the most aggressive behaviors most often while G. assimilis transitioned between two of the least aggressive behaviors most often. We integrate this new information on aggression for G. assimilis and G. veletis with previously documented aggression data for many cricket species to investigate aggression in a broader evolutionary context than previously possible. Within a phylogenetic context, we test the hypothesis that species whose males use burrows from which to call and attract females are more aggressive than species with non‐burrowing males. We found evidence consistent with this hypothesis; species with burrowing males tended to be more aggressive than species with non‐burrowing males. Together, our study provides fine‐scale understanding of aggression in two cricket species and broad‐scale evolutionary context for aggression across cricket species.     

Asymmetric Forceps Increase Fighting Success among Males of Similar size in the Maritime Earwig

Extreme asymmetric morphologies are hypothesized to serve an adaptive function that counteracts sexual selection for symmetry. However, direct tests of function for asymmetries are lacking, particularly in the context of animal weapons. The weapon of the maritime earwig, Anisolabis maritima, exhibits sizeable variation in the extent of directional asymmetry within and across body sizes, making it an ideal candidate for investigating the function of asymmetry. In this study, we characterized the extent of weapon asymmetry, characterized the manner in which asymmetric weapons are used in contests, staged dyadic contests between males of different size classes, and analyzed the correlates of fighting success. In contests between large males, larger individuals won more fights and emerged as the dominant male. In contests between small males, however, weapon asymmetry was more influential in predicting overall fighting success than body size. This result reveals an advantage of asymmetric weaponry among males that are below the mean size in the population. A forceps manipulation experiment suggests that asymmetry may be an indirect correlate of a morphologically independent factor that affects fighting ability.     

Size-biased Mating in Both Sexes of the Black-horned Tree Cricket, <Emphasis Type="Italic">Oecanthus nigricornis</Emphasis> Walker (Orthoptera: Gryllidae: Oecanthinae)

Previous studies on tree crickets have demonstrated female choice of males based on size and courtship feeding but less is known about sexual selection under conditions of direct mating competition. I studied courtship, aggression and mating of the black-horned tree cricket Oecanthus nigricornis (Walker) to test size-related sexual selection under conditions of direct sexual competition. Results show that larger individuals of both sexes mated more frequently than their smaller counterparts, and this was due to the ability of large individuals to out compete rivals. Large males achieved the advantage by aggressively reducing courtship by small males, whereas large females responded to male courtship more quickly but with little aggression. Although there was no evidence here for mate choice, there were advantages for having larger mates; fecundity increased with female size and spermatophores (which females consume after mating) increased with male size. Size of the specialized metanotal courtship gift, however, was not related to male size.     

Weaponry, color, and contest success in the jumping spider Lyssomanes viridis

Weaponry and color badges are commonly theorized to function as visual signals of aggressiveness or fighting ability. However, few studies have supported a signaling function of weaponry, and the role of color in invertebrate competitive interactions remains virtually unexplored. Jumping spiders (Salticidae) make excellent invertebrate models for studying weaponry and color because males of many species are colorful and possess exaggerated chelicerae, which are used as weapons in escalated contests. To determine whether color or weaponry might function as visual signals in male-male competitions, we investigated relationships between contest success, cheliceral length, and red coloration in Lyssomanes viridis. Males having longer chelicerae than their opponents were significantly more likely to win (p=0.0008). Males who won, despite being smaller than their opponents, had significantly less red chelicerae than their opponents (p=0.01). Male and female cheliceral length, as well as foreleg length, correlated tightly with body size. Cheliceral and foreleg length showed significantly stronger positive allometry in males than in females. We conclude that male chelicerae and forelegs are under strong positive selection for their use in physical fights and/or as visual signals of fighting ability.     

Calling,Courtship, and Condition in the Fall Field Cricket,Gryllus pennsylvanicus

Theoretically, sexual signals should provide honest information about mating benefits and many sexually reproducing species use honest signals when signalling to potential mates. Male crickets produce two types of acoustic mating signals: a long-distance mate attraction call and a short-range courtship call. We tested whether wild-caught fall field cricket (Gryllus pennsylvanicus) males in high condition (high residual mass or large body size) produce higher effort calls (in support of the honest signalling hypothesis). We also tested an alternative hypothesis, whether low condition males produce higher effort calls (in support of the terminal investment hypothesis). Several components of long-distance mate attraction calls honestly reflected male body size, with larger males producing louder mate attraction calls at lower carrier frequencies. Long-distance mate attraction chirp rate dishonestly signalled body size, with small males producing faster chirp rates. Short-range courtship calls dishonestly reflected male residual mass, as chirp rate and pulse rate were best explained by a curvilinear function of residual mass. By producing long-distance mate attraction calls and courtship calls with similar or higher effort compared to high condition males, low condition males (low residual mass or small body size) may increase their effort in current reproductive success at the expense of their future reproductive success, suggesting that not all sexual signals are honest.     

Adaptive Plasticity in Wild Field Cricket’s Acoustic Signaling

Phenotypic plasticity can be adaptive when phenotypes are closely matched to changes in the environment. In crickets, rhythmic fluctuations in the biotic and abiotic environment regularly result in diel rhythms in density of sexually active individuals. Given that density strongly influences the intensity of sexual selection, we asked whether crickets exhibit plasticity in signaling behavior that aligns with these rhythmic fluctuations in the socio-sexual environment. We quantified the acoustic mate signaling behavior of wild-caught males of two cricket species, Gryllus veletis and G. pennsylvanicus. Crickets exhibited phenotypically plastic mate signaling behavior, with most males signaling more often and more attractively during the times of day when mating activity is highest in the wild. Most male G. pennsylvanicus chirped more often and louder, with shorter interpulse durations, pulse periods, chirp durations, and interchirp durations, and at slightly higher carrier frequencies during the time of the day that mating activity is highest in the wild. Similarly, most male G. veletis chirped more often, with more pulses per chirp, longer interpulse durations, pulse periods, and chirp durations, shorter interchirp durations, and at lower carrier frequencies during the time of peak mating activity in the wild. Among-male variation in signaling plasticity was high, with some males signaling in an apparently maladaptive manner. Body size explained some of the among-male variation in G. pennsylvanicus plasticity but not G. veletis plasticity. Overall, our findings suggest that crickets exhibit phenotypically plastic mate attraction signals that closely match the fluctuating socio-sexual context they experience.     

Reproductive Success and Body Size in the Cricket Gryllus firmus

Male body size influences mate choice and sexual selection in many animal species. Here we investigate the role of male body size in the reproductive success of the field cricket Gryllus firmus. This species hybridizes with a close smaller relative, G. pennsylvanicus, and it is thought that this size difference may affect reproductive isolation between these species. We paired large and small G. firmus males with a single G. firmus female and genotyped the resulting offspring. Overall, larger males sired a greater proportion of offspring and in a majority of the crosses the larger male sired all of the offspring. For crosses in which both males sired offspring, there was no difference in the proportion of offspring sired by small and large males. Intrasexual competition, female choice, and differences in ejaculates between males could all influence the patterns we observe. We discuss the implications of our findings within the context of reproductive isolation between G. firmus and G. pennsylvanicus.     

Territory Availability at Different Spatial Scales does not Affect Fighting Investment in Males of the Lycaenid Butterfly Strymon mulucha

In territorial species, rivals investment in fights over territories may increase when the availability of suitable areas for defense is low. This should occur because low territory availability may increase the costs to maintain and acquire territories. Although such process occurs in small spatial scales (local scale), territory availability in larger scales (regional scale) may also affect fighting investment, as losers should incur additional dispersing costs to find new territories. In this study, we used males of the hilltopping butterfly Strymon mulucha to evaluate the hypothesis that males should invest more in territorial fights when the costs to find new territories are higher (both at local and at regional scale). We timed male–male contests for territories located in 12 hilltops and measured male density per territory in each hilltop (local scale). We also quantified the distance between hilltops containing suitable areas for territories (regional scale). Male–male contests lasted 21 s on average, and copulations did not occur during the observations. The duration of contests was unrelated to the male density per territory or to the distance among hilltops, indicating that the investment in fights was unaffected by the availability of territorial sites, independent of the spatial scale. As male–male contests in S. mulucha are longer than the mean contest duration in other butterfly species and mating is extremely rare, we suggest that the value of each territory may be high enough to favor males that always invest as much as possible in contests.     

Overt and covert competition in a promiscuous mammal: the importance of weaponry and testes size to male reproductive success

Male contests for access to receptive females are thought to have selected for the larger male body size and conspicuous weaponry frequently observed in mammalian species. However, when females copulate with multiple males within an oestrus, male reproductive success is a function of both pre- and postcopulatory strategies. The relative importance of these overt and covert forms of sexual competition has rarely been assessed in wild populations. The Soay sheep mating system is characterized by male contests for mating opportunities and high female promiscuity. We find that greater horn length, body size and good condition each independently influence a male's ability to monopolize receptive females. For males with large horns at least, this behavioural success translates into greater siring success. Consistent with sperm-competition theory, we also find that larger testes are independently associated with both higher copulation rates and increased siring success. This advantage of larger testes emerges, and strengthens, as the number of oestrous females increases, as dominant males can no longer control access to them all. Our results thus provide direct quantitative evidence that male reproductive success in wild populations of mammals is dependent upon the relative magnitude of both overt contest competition and covert sperm competition.     

Imperfect assessment and limited information preclude optimal strategies in male-male fights in the orb-weaving spider Metellina mengei

Agonistic behaviour between male orb-web spiders Metellina mengei competing for access to female webs was examined in field experiments to test the major predictions of game theory. Winners of fights were significantly larger than losers, particularly with respect to the length of the first pair of legs, which are sexually dimorphic in this species and used extensively in agonistic encounters. The size of the winning male had no influence on contest intensity or duration, and neither did relative size. However, fight intensity and duration were both positively correlated with the size of the losing male. Resident males won significantly more contests than intruders. Winning intruders were significantly larger than winning residents and it was these winning intruders that tended to produce the longer fights. Female weight and hence reproductive value had a marked influence on fight intensity and duration of fights won by the intruder but not those won by the resident. This indicates that only the resident obtains information about the female. These data are discussed with reference to the discrepancy with theory and a failure of some contestants to obtain information on resource value and relative contestant size necessary to optimize fight strategy.     

The larval head of Exechia (Mycetophilidae) and Bibio (Bibionidae) (Diptera)

Exechia and Bibio have retained several plesiomorphic groundplan features of Diptera and Bibionomorpha, including a fully exposed and sclerotized head capsule, the transverse undivided labrum, the absence of movable premandibles, and undivided mandibles without combs. The fusion of the hypostomal bridge with the head capsule and largely reduced antennae are derived features shared by both taxa. The absence of teeth at the anterior hypostomal margin is a potential autapomorphy of Bibionomorpha. A basal position of Anisopodidae is suggested by a number of plesiomorphies retained in this family. Apomorphies of Bibionomorpha excluding Anisopodidae are the reduction of tentorial elements, the partial fusion of the labrum and clypeus, one-segmented antennae, the absence of a separate submental sclerite, the loss of the labial palpus, and the reduction of the pharyngeal filter apparatus. Head structures of Bibio are largely unmodified. The subprognathous orientation is one of few autapomorphic features. In contrast, the mouthparts of Exechia are highly modified in correlation with the specialized food uptake. The rasping counterrotating movements of maxillae and mandibles with teeth oriented in opposite directions are carried out by strongly developed extensors and flexors of the paired mouthparts. The modified labium mechanically supports the “drill head” formed by the mandibles und maxillae. The necessary stability of the head capsule is provided by the hypostomal bridge which also compensates the far-reaching reduction of the tentorium.     

Sex and Weapons: Contrasting Sexual Dimorphisms in Weaponry and Aggression in Snapping Shrimp

Sexual dimorphisms in weaponry and aggression are common in species in which one sex (usually males) competes for access to mates or resources necessary for reproduction – sexually dimorphic weaponry and aggression, in other words, are frequently the result of intrasexual selection. In snapping shrimp, the major chela (snapping claw) can be a deadly weapon, and males of many species have larger chelae than females, a pattern readily interpreted as resulting from intrasexual selection. Thus, males might be expected to show more sex‐specific aggression than females, and be more aggressive overall. We tested these predictions in two species of snapping shrimp in a territorial defense context. Neither of these predictions was supported: in both species, females, but not males, engaged in sex‐specific aggression and females were more aggressive than males overall. These contrasting sexual dimorphisms – larger weaponry in males but higher aggression in females – highlight the importance of considering the function of weaponry and aggression in contexts other than direct competitions over mates. In addition, species differences in the degree of sexual dimorphism in chela size were due to differences in female, not male, chela size, and the species with greater sexual dimorphism in weaponry was significantly less aggressive overall; also, while paired and solitary males did not differ in residual chela size, for the species with greater sexual dimorphism, females carrying embryos had smaller residual chela sizes. These results suggest that understanding the sexual dimorphisms in weaponry and aggression in snapping shrimp requires understanding the relative costs and benefits of both in females as well as males.     

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

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

Relationships between body size and sound‐producing structures in crickets: do large males have large harps?

Abstract. Male crickets produce conspicuous acoustic signals to attract mates and deter potential rivals. These signals are created when a male cricket closes his wings rapidly and a file and scraper mechanism causes several areas of the wing to vibrate. The harp is an area of the wing that is part of the resonating structure. Because the harp acts as part of a mechanical resonator, changes in harp area or mass could influence the frequency of sound produced. Because females exhibit stabilizing selection on the frequency used in male songs, we hypothesized that there would be a negative allometric relationship between body size and harp area. In addition, we examined the degree of asymmetry in the harp, wing, and tibia. We examined this in four different species of cricket: Acheta domesticus, Gryllus bimaculatus, Gryllus rubens , and Teleogryllus oceanicus. For each species, we measured pronotum width as an index of body size, tibia length, and the area of the forewing and harp. There were significant differences among species in their morphological characteristics. We observed consistent directional asymmetry in the harp area but not in the total wing area. When wings did exhibit directional asymmetry, it was in the opposite direction of the directional asymmetry observed in the harp. Within species, larger males typically had larger harps and the relationship between harp area and body size exhibited negative allometry. Wing area exhibited an isometric relationship with body size. Our data provide a potential mechanism linking decreases in song frequency with body size in male crickets, and suggest that sensory constraints might influence the morphology of signaling structures in a similar fashion as genitalia.     

Dominance is not always an honest signal of male quality,but females may be able to detect the dishonesty

Females prefer dominant males as mating partners in numerous species. Male dominance rank is considered as an honest signal of male quality, because only healthy males in good condition are thought to be able to win fights with other males. Here, we tested whether activation of the immune system influences the success of males in male–male competition and mating in the field cricket, Gryllus integer. We activated the immune system of males with a nylon monofilament (to mimic a parasitoid larva), and arranged fights between male pairs to assess male dominance and associated mating success. Activation of the immune system with nylon monofilament substantially enhanced the fighting success of males during male–male competition but had no effect on mating success. However, sham-manipulation (a wound only) did not have any effect on fighting success although females mated more often with dominant males. Our study suggests that when male crickets meet an apparent survival threat they may behave more dominantly, probably owing to terminal investment. Male success during male–male competition is not always an honest signal of males’ quality, but females may be able to detect this dishonesty.     

Removal of both antennae influences the courtship and aggressive behaviors in male crickets

To test whether insect antennae are necessary for eliciting courtship and aggression toward appropriate partners, we antennectomized adult male crickets (Gryllus bimaculatus) and observed their behavior toward other antennectomized males and intact females. At 7 days after removal of both antennae, pairs of antennectomized males were placed together; 80% displayed courtship behavior, generating courtship song by rubbing their forewings together, toward other antennectomized males, and 20% displayed aggressive behavior. Only 45% courted intact females. No intact males courted antennectomized males, and 80% displayed aggressive behavior. All intact males courted females. The results for males with one antenna removed were essentially the same as for intact males. These findings indicate that a high proportion of male crickets with both antennae removed court other males and fail to display male-male aggression, demonstrating that removal of antennae from male crickets induces male-male courtship and that an antenna is necessary for the expression of male-male aggression. Moreover, brain serotonin (5-hydroxytryptamine; 5-HT) levels in male crickets were significantly reduced at 7 days after removal of antennae. The reduction of 5-HT was detected primarily in the central body of the brain. Thus, 5-HT in the central body of the male cricket brain may be involved in the behavioral changes.     

UV-Green Iridescence Predicts Male Quality during Jumping Spider Contests

Animal colour signals used in intraspecies communications can generally be attributed to a composite effect of structural and pigmentary colours. Notably, the functional role of iridescent coloration that is ‘purely’ structural (i.e., absence of pigments) is poorly understood. Recent studies reveal that iridescent colorations can reliably indicate individual quality, but evidence of iridescence as a pure structural coloration indicative of male quality during contests and relating to an individual’s resource-holding potential (RHP) is lacking. In age- and size-controlled pairwise male-male contests that escalate from visual displays of aggression to more costly physical fights, we demonstrate that the ultraviolet-green iridescence of Cosmophasis umbratica predicts individual persistence and relates to RHP. Contest initiating males exhibited significantly narrower carapace band separation (i.e., relative spectral positions of UV and green hues) than non-initiators. Asymmetries in carapace and abdomen brightness influenced overall contest duration and escalation. As losers retreated upon having reached their own persistence limits in contests that escalated to physical fights, losers with narrower carapace band separation were significantly more persistence. We propose that the carapace UV-green iridescence of C. umbratica predicts individual persistence and is indicative of a male’s RHP. As the observed UV-green hues of C. umbratica are ‘pure’ optical products of a multilayer reflector system, we suggest that intrasexual variations in the optical properties of the scales’ chitin-air-chitin microstructures are responsible for the observed differences in carapace band separations.