Male Weapon Research Articles

Size as a determinant of robustness: Exploring sexual dimorphism in the size and shape of the snapping and cutting claws in Alpheus (caridea: Alpheidae)

This study investigates the role of static allometry in modulating sexual shape dimorphism in the snapping and cutting claws of Alpheus species. We hypothesize that the static allometry in adulthood can promote an exaggeration of existing sexual dimorphism in the snapping and cutting claws of Alpheus. Sexually mature pairs from three species (Alpheus angulosus, Alpheus carlae, and Alpheus nuttingi) were analyzed using geometric morphometric techniques. The increase in size claw enhances the robustness of both snapping and cutting claws. Since males carry larger weapons compared to females, the static allometry provides more robustness in their weapons. These findings support our hypothesis. Due to the effect of static allometry, male weapons exhibit a more robust pollex, while female weapons exhibit a more robust palm in the snapping claw. Sexual shape dimorphism is less pronounced in the cutting claw, although males display larger and more robust claws compared to females. Static allometry amplifies the sexual differences in Alpheus weapons, emphasizing the role of sexual selection in the evolution of claw morphology. The results suggest that sexual selection favors larger, more robust claws in males, thus enhancing their effectiveness in agonistic contests.

Energetically costly weaponry in the large morph of male stag beetles

AbstractMales of many species develop disproportionately large weapons, and frequently the weapon size increases more quickly with body size in small than in large males—a trend often considered to result from the depletion of developmental resources limiting male weapon exaggeration. Based on the cost‐minimization hypothesis, a shallower weapon‐body size allometric slope in large than in small males could be selected for if the energetic maintenance costs of having oversized weapons increase faster in large than in small males. Whether males of different size groups that differ in the weapon‐body size allometric relationship also differ in the energetic costs of maintaining their weapons has not yet been investigated. Consequently, we studied this using the stag beetle (Cyclommatus mniszechi), in which males are larger in size and have larger mandibles than do females and are divided into two morphs: majors are larger and have longer mandibles than minors. In both male morphs, mandible size increases disproportionately with their body size, but it increases more quickly in minors than in majors. We measured the standard metabolic rate (SMR) of the three types of these beetles (the two male morphs and females) to investigate whether the energetic maintenance cost of carrying weapons differs among them. Body weight explained 80% of the variation in the SMR in the three types of beetles, and SMR increased more quickly with body weight in major males than in minor males or females. Weapon and body size also jointly explained approximately 80% of the variation in the SMR. Whereas SMR increased with body size in all three types of beetles (with a higher rate of increase in major males than in minor males or females), only the major males' SMR increased with weapon size. Overall, being heavier and larger and carrying oversized weapons are energetically costly in major males, something which could constrain the exaggeration of their weapons.

Effect of temperature on sexual size dimorphism during the developmental period in the broad-horned flour beetle

Adult size in numerous insects is strongly dependent on temperature. In several cases, a temperature–size rule is observed in which developmental temperature and adult size tradeoff. Although several previous studies have demonstrated the temperature–size rule, only a few have explored the relationship between developmental temperature and weapon traits or sexual size dimorphism. This study was conducted to investigate the size of the broad-horned flour beetle Gnatocerus cornutus when it was developed under different temperatures. G. cornutus males possess weapon traits for male–male combat and exhibit sexual size dimorphism in other morphological traits. Results showed that male weapon size and body size complied with the temperature–size rule. Furthermore, the extent of sex dimorphism in genae width, a weapon-supportive trait, were larger at lower temperatures. Our findings suggest that the temperature–size rule also influences the size of sexual traits.

Extreme range in adult body size reveals hidden trade-offs among sexually selected traits.

Sexually selected weapons used to monopolize mating opportunities are predicted to trade-off with traits used in competition for fertilization. Yet, the limited size range typically found among adults of a species often precludes clear comparisons between population-level and individual-level relative trait investment. The jousting weevil, Brentus anchorago (Coleoptera: Brentidae), varies more than 26-fold in body mass, which is among the most extreme adult body size ranges of any solitary terrestrial species. We reveal a trade-off at a population level: hypermetric scaling in male weapons (slope = 1.59) and a closely mirrored reversal in allocation to postcopulatory traits (slope = 0.54). Yet, at the individual level, we find the opposite pattern; males that invest relatively more in weapons for their size class also invest more in postcopulatory traits. Across 36 dung beetle and 41 brentine weevil species, we find the allometric slope explains more trait variation at larger body size ranges; in brentines, population-level scaling patterns become more detectable in species with a larger range in adult body size. Our findings reveal that population-level allometries and individual-level trade-offs can both be important in shaping relative trait allocation; we highlight that the adult body size range is rarely examined but may be integral to gaining a deeper understanding of trade-offs in reproductive allocation.

Alternative mating strategies in the Wellington tree weta expressed via genetic polymorphism for precocial male maturation.

The expression of alternative mating strategies is generally hypothesized to be a plastic response of the genome to the environment or a genetic polymorphism. The Wellington tree weta (Hemideina crassidens) is an orthopteran insect in which males express three alternative mating morphotypes related to the size of mandibular weaponry. My common garden experiment in the laboratory shows that male weapon size at maturity in this species is nether condition- nor environment-dependent; therefore, supporting the hypothesis that it is genetically polymorphic. Males that matured at the 8th instar had the shortest development time and the smallest weaponry, males that matured at the 10th instar had the longest development time and the largest weaponry, and males that matured at the 9th instar were intermediate to the other two morphs. These morph-related differences in development time suggest that male alternative mating strategies in this species are expressed via precocial male maturation, a rare mode of strategy expression in insects. Additionally, adult lifespans differed significantly among morphs with 8th instar males having the shortest, and 9th instar males having the longest lifespans. I discuss how these differences in lifespan align with morph-specific investment in pre- and post-copulatory processes to equalize fitnesses among morphs.

Inbreeding depression in a sexually selected weapon and the homologue in females.

Theory predicts that traits with heightened condition dependence, such as sexually selected traits, should be affected by inbreeding to a greater degree than other traits. The presence of environmental stress may compound the negative consequences of inbreeding depression. In this study, we examined inbreeding depression across multiple traits and whether it increased with a known form of environmental stress. We conducted our experiment using both sexes of the sexually dimorphic leaf-footed cactus bug, Narnia femorata (Hemiptera: Coreidae). Adult male cactus bugs have enlarged hind legs used as weapons in male-male contests; these traits, and their homologue in females, have been previously found to exhibit high condition dependence. In this study, we employed a small developmental group size as an environmental stress challenge. Nymph N. femorata aggregate throughout their juvenile stages, and previous work has shown the negative effects of small group size on survivorship and body size. We found evidence of inbreeding depression for survival and seven of the eight morphological traits measured in both sexes. Inbreeding depression was higher for the size of the male weapon and the female homolog. Additionally, small developmental group size negatively affected survival to adulthood. However, small group size did not magnify the effects of inbreeding on morphological traits. These findings support the hypothesis that traits with heightened condition dependence exhibit higher levels of inbreeding depression.

Sexually-selected male weapon increases the risk of population extinction under environmental change: an experimental evidence.

Exaggerated sexually-selected traits, occurring more commonly in males, help individuals to increase reproductive success, but are costly to produce and maintain. These costs on the one hand may improve population fitness by intensifying selection against maladapted males, but on the other hand may increase the risk of extinction under environmental challenge. However, the impact of sexually selected traits on extinction risk have not been investigated experimentally. We used replicate populations of a male-dimorphic mite, Rhizoglyphus robini, to test if prevalence of a sexually-selected weapon affected the risk of extinction under temperature increase (2 0C per each of three consecutive generations). In two independent experiments that utilized either inbred lines or lines mass selected for or against the weapon to establish experimental replicate populations differing in the prevalence of the weapon, we found that populations with high weapon prevalence were more likely to go extinct. Extinctions occurred despite partial suppression of the weapon expression at increased temperature and were not explained by increased male mortality. Our results provide the first, to our knowledge, experimental evidence demonstrating dramatic effect of elaborated sexual traits on the risk of extinction under environmental challenge.

A sexually selected male weapon characterized by strong additive genetic variance and no evidence for sexually antagonistic polyphenic maintenance.

Sexual selection and sexual antagonism are important drivers of eco-evolutionary processes. The evolution of traits shaped by these processes depends on their genetic architecture, which remains poorly studied. Here, implementing a quantitative genetics approach using diallel crosses of the bulb mite, Rhizoglyphus robini, we investigated the genetic variance that underlies a sexually selected weapon that is dimorphic among males and female fecundity. Previous studies indicated that a negative genetic correlation between these two traits likely exists. We found male morph showed considerable additive genetic variance, which is unlikely to be explained solely by mutation-selection balance, indicating the likely presence of large-effect loci. However, a significant magnitude of inbreeding depression also indicates that morph expression is likely to be condition-dependent to some degree and that deleterious recessives can simultaneously contribute to morph expression. Female fecundity also showed a high degree of inbreeding depression, but the variance in female fecundity was mostly explained by epistatic effects, with very little contribution from additive effects. We found no significant genetic correlation, nor any evidence for dominance reversal, between male morph and female fecundity. The complex genetic architecture underlying male morph and female fecundity in this system has important implications for our understanding of the evolutionary interplay between purifying selection and sexually antagonistic selection.

Do reproductive characteristics of the species explain differences in the investment in weapon size present in males?

AbstractSexually selected traits, such as male weapons, are highly variable in shape and size across species. However, little is known about the mechanisms that may govern this variation. Because males with greater investment in weapon size tend to win more fights, but also pay higher costs related to weapon development and maintenance, larger weapons should be expected only in species in which victory in male–male fights generate reproductive benefits that outweigh investment costs. Here, we hypothesized that the reproductive characteristics that increase the chances of winners to access females or to fertilize eggs will favor the evolution of larger weapons. To evaluate this hypothesis, we conducted a meta‐analysis using arthropods as model organisms. To measure investment in weapon size, we gathered both Pearson correlation coefficient and the standardized (but non centralized) slope values for the relationship between weapon size and body size. We found that none of the reproductive characteristics we investigated was related to male weapon size. Thus, it seems that greater certainty of accessing a female or fertilizing female eggs with a victory does not modulate the investment in male weapon size. Perhaps the cost–benefit ratio between weapon size investment and reproductive success is not the main factor driving the variation in weapon size.

Functional roles of multiple male weapons in the flower beetle Dicronocephalus wallichii

Functional roles of multiple male weapons in the flower beetle <i>Dicronocephalus wallichii</i>

Biomechanics influence sexual dimorphism in the giant mesquite bug, Thasus neocalifornicus

In many species, males possess specialized weaponry that confers benefits during male-male combat. Because male weapons are often disproportionately larger versions of preexisting body parts, females often possess reduced versions of male weaponry. Most research focuses exclusively on sexual dimorphism in the size of male and female weapons, even though other aspects such as weapon performance can also explain the evolution of weapon sexual dimorphism. In the giant mesquite bug, Thasus neocalifornicus, males wield exaggerated hindlegs that aid in locomotion and are used as weapons to generate forceful squeezes during combat. However, female T. neocalifornicus hindlegs are relatively inconspicuous and only used for locomotion. To understand the intricacies of weapon sexual dimorphism in T. neocalifornicus hindlegs, we measured the allometry of their hindlegs morphology, biomechanics, and performance. Males and females had relatively similar sized legs when concerning only linear measurements: hindleg length did not differ between the sexes (both for intercept and slope), but males do have relatively wider hindlegs (greater intercepts). Regarding performance, however, males were relatively and proportionally stronger than females. Furthermore, the output lever of male hindlegs scales hypoallometrically and the tibial spine maintains its position as the hindlegs grows, both of which maintain the hindlegs’ biomechanical efficiency as they increase in size. Overall, our finding demonstrates that selection on the performance and biomechanics of sexually selected weapons can influence the expression of sexual dimorphism, by exaggerating some aspects of the weapons morphology—but constraining others.

It is not always about body size: evidence of Rensch's rule in a male weapon.

In many species, sexual dimorphism increases with body size when males are the larger sex but decreases when females are the larger sex, a macro-evolutionary pattern known as Rensch's rule (RR). Although empirical studies usually focus exclusively on body size, Rensch's original proposal included sexual differences in other traits, such as ornaments and weapons. Here, we used a clade of harvestmen to investigate whether two traits follow RR: body size and length of the fourth pair of legs (legs IV), which are used as weapons in male-male fights. We found that males were slightly smaller than females and body size did not follow RR, whereas legs IV were much longer in males and followed RR. We propose that sexual selection might be stronger on legs IV length than on body size in males, and we discuss the potential role of condition dependence in the emergence of RR.

Digest: Sexually selected weapons: Winning the fight, but losing the war for reproduction on a changing battlefield*

What conditions favor the evolution of large animal weapons? In the Japanese rhinoceros beetle, Trypoxylus dichotomus, del Sol et al. found that selection favors large horns in populations where males compete over guardianship of scarce female feeding territories. However, in other populations, an abundance of female feeding territories reduces the chance of mating success for these guarding males, leading to the evolution of relatively shorter horn sizes. This finding illustrates that female habitat and resource use have the potential to influence evolution of male weapon size through sexual selection.

Anti-predator behaviour depends on male weapon size.

Tonic immobility and escape are adaptive anti-predator tactics used by many animals. Escape requires movement, whereas tonic immobility does not. If anti-predator tactics relate to weapon size, males with larger weapons may adopt tonic immobility, whereas males with smaller weapons may adopt escape. However, no study has investigated the relationship between weapon size and anti-predator tactics. In this study, we investigated the relationship between male weapon size and tonic immobility in the beetle Gnathocerus cornutus. The results showed that tonic immobility was more frequent in males with larger weapons. Although most studies of tonic immobility in beetles have focused on the duration, rather than the frequency, tonic immobility duration was not affected by weapon size in G. cornutus. Therefore, this study is the first, to our knowledge, to suggest that the male weapon trait affects anti-predator tactics.

Harder, better, faster, stronger: Weapon size is more sexually dimorphic than weapon biomechanical components in two freshwater anomuran species.

Sexual selection influences the evolution of morphological traits that increase the likelihood of monopolizing scarce resources. When such traits are used during contests, they are termed weapons. Given that resources are typically linked to monopolizing mating partners, theory expects only males to bear weapons. In some species, however, females also bear weapons, although typically smaller than male weapons. Understanding why females bear smaller weapons can thus help us understand the selective pressures behind weapon evolution. However, most of our knowledge comes from studies on weapon size, while the biomechanics of weapons, such as the size of the muscles, efficiency, and shape are seldom studied. Our goal was to test if the theoretical expectations for weapon size sexual dimorphism also occur for weapon biomechanics using two aeglid crab species. Males of both species had larger claws which were also stronger than female claws. Male claws were also more efficient than females' claws (although we used only one species in this analysis). For weapon shape, though, only one species differed in the mean claw shape. Regarding scaling differences, in both species, male claws had higher size scaling than females, while only one species had a higher shape scaling. However, male weapons did not have higher scaling regarding strength and efficiency than females. Thus, males apparently allocate more resources in weapons than females, but once allocated, muscle and efficiency follow a similar developmental pathway in both sexes. Taken together, our results show that sexual dimorphism in weapons involves more than differences in size. Shape differences are especially intriguing because we cannot fully understand its causes. Yet, we highlight that such subtle differences can only be detected by measuring and analysing weapon shape and biomechanical components. Only then we might better understand how weapons are forged.

«AND GIRLS AMAZONS ACCUSTOMED TO ARMS, HORSES AND HUNTING…»

According to ancient authors Amazons treated their new-borns differently. They usually gave the boys to their fathers. At the same time, they raised the girls themselves. Moreover, the girls since childhood were taught martial art and hunting. That is, first of all they were taught archery and horseback riding. Based on these data, we drew attention to the Scythian girls’ graves with weapons.&#x0D; 17 such burials of girls are known today in the area of European Scythia. They are fixed on the territory of the Steppe Dnieper Region, Crimea, Transnistria and Don Region.&#x0D; All children’s graves are individual except for one (with a teenager-servant). Only two of them were robbed in antiquity.&#x0D; All graves (with one exception) were excavated in the kurhans. An analysis of the types of burial structures showed that undercut graves and catacombs predominate. Simple pits are represented by only two cases. The spread of these types of graves on different regions coincides with the same graves of adult Amazons.&#x0D; All burials were made according to the rite of inhumation. The western orientation of the dead is predominant. In general, burial sites, rites and anthropological types of Amazons are identical to those recorded for the population of European Scythia.&#x0D; Material support is divided in three main groups: 1) universal (meat meals with a knife and utensils); 2) Female (spindles with spinners, earrings, necklace, bracelets, rings, stone dishes); 3) Male (weapons, horse bridles, hryvnias). The combination of the components of these kits has usually marked the burial of the Amazons. The nature of the weapon shows that it was intended primarily for the remote combat.&#x0D; The evidence of social stratification is, first of all, the size of burial structures and the composition of material support. According to these indicators, three social groups are distinguished in the Amazon girls’ environment.&#x0D; The chronological range of the burial complexes of girls with weapons is determined from the end of the 5th to 3—2 centuries BC. However, the main group of these tombs is dated 4th century B.C.

Defining an intrasexual male weapon polymorphism in a New Zealand harvestman (Opiliones: Neopilionidae) using traditional and geometric morphometrics

Abstract In many species, competition for mates has led to exaggerated male sexually-selected traits. Sexually-selected male weapons are used in male-male combat and include structures like horns, antlers and enlarged teeth. Weapons often vary intraspecifically in size, resulting in either a continuum of weapon sizes or in discrete male polymorphisms. More rarely, complex weapon polymorphisms can also include variation in weapon shape; however, these are difficult to quantify. Here we first use traditional linear morphometrics to describe a weapon trimorphism in the endemic New Zealand harvestman, Forsteropsalis pureoraTaylor, 2013. We identified three male morphs: a small-bodied gamma male with reduced chelicera, a large-bodied beta male with long, slender chelicerae, and a large-bodied alpha male with shorter, but very broad, robust chelicerae. Chelicera length alone failed to fully capture the variation in weapon investment. Using geometric morphometrics, we show that alpha males are different in weapon shape, whereas beta and gamma males have similar weapon shape, but vary in their body size and chelicera length. Additionally, we describe how the chelicerae function during male-male combat from observations of contests. This work demonstrates how combining linear and geometric morphometrics can help to elucidate complex polymorphism.

Selection for Male Weapons Boosts Female Fecundity, Eliminating Sexual Conflict in the Bulb Mite

Extreme differences between the sexes are usually explained by intense sexual selection on male weapons or ornaments. Sexually antagonistic genes, with a positive effect on male traits but a negative effect on female fitness, create a negative inter-sexual correlation for fitness (sexual conflict). However, such antagonism might not be apparent if sexually selected male traits are condition-dependent, and condition elevates female fitness. Here we reveal a surprising positive genetic correlation between male weaponry and female fecundity. Using mite lines that had previously been through 13 generations of selection on male weapons (fighting legs), we investigated correlated evolution in female fecundity. Females from lines under positive selection for weapons (up lines) evolved higher fecundity, despite evolving costly, thicker legs. This is likely because male mites have condition-dependent weaponry that increases our ability to indirectly select on male condition. Alleles with positive effects on condition in both sexes could have generated this correlation because: the up lines evolved a higher proportion of fighters and there were positive correlations between weapon size and the male morph and sex ratios of the offspring. This positive inter-sexual genetic correlation should boost the evolution of male weapons and extreme sex differences.

Pay attention to the ladies: female aggressive behavior and weapon allometry provide clues for sexual selection in freshwater anomurans (Decapoda: Aeglidae)

Contesting scarce resources can trigger the evolution of specialized morphological structures (i.e., animal weapons). While most research focus on male weapons, females might also bear weapons, although generally smaller and less conspicuous than male weapons. Social selection is evoked to explain female weaponry in which females fight for nonsexual resources such as food and shelter. Males might fight for similar resources but are expected to have proportionally larger weapons due to additional inputs from sexual selection. We tested whether males have proportionally larger weapons than females in two species of Aegla crabs. Interestingly, only males of one species had proportionally larger claws than females. Given that these larger weapons typically correlate to increased aggression, we expected males to fight more intensely than females. Thus, we compared intrasexual contests of males and females of the same species. Females fought similarly to males: latency, contest duration, and frequency of highly aggressive acts were similar between the sexes. Therefore, despite males having proportionally larger weapons in one species (as predicted by sexual selection), they fought similarly to females. Our results hint that fighting might not necessarily be the source of selection for sexual dimorphism we typically expect. Other sources, such as the frequency of fighting and predation pressure, might be selecting larger claws in males despite the similar fights, while fecundity costs might downsize female claws. We highlight that comparing female with male weapons and the associated fighting behavior shows that selection on weapons is not as straightforward as we might think. We show that studying the highly neglected female weapon allometry and usage allows us to infer on the selective process acting on animal weapons. We show here that males of one species have proportionally larger weapons, and, despite that, males fight similarly to females in every aspect analyzed. Therefore, fighting per se might not be the sole source of selection to explain sexual dimorphism. Other sources, such as frequency of fighting, and how expensive it is to produce gametes might be additional sources of selection.

Fluctuating selection strength and intense male competition underlie variation and exaggeration of a water strider's male weapon.

Sexually selected traits can reach high degrees of phenotypic expression and variation under directional selection. A growing number of studies suggest that such selection can vary in space, time and form within and between populations. However, the impact of these fluctuations on sexual trait evolution is poorly understood. In the water strider Microvelia longipes, males display striking trait exaggeration and phenotypic variation manifested as extreme differences in the rear leg length. To study the origin and maintenance of this exaggerated trait, we conducted comparative behavioural, morphometric and reaction norm experiments in a selection of Microvelia species. We uncovered differences both in the mating behaviour and the degree of sexual dimorphism across these species. Interestingly, M. longipes evolved a specific mating behaviour where males compete for egg-laying sites, consisting of small floating objects, to intercept and copulate with gravid females. Through male–male competition assays, we demonstrated that male rear legs are used as weapons to dominate egg-laying sites and that intense competition is associated with the evolution of rear leg length exaggeration. Field observations revealed rapid fluctuation in M. longipes habitat stability and the abundance of egg-laying sites. Paternity tests using genetic markers demonstrated that small males could only fertilize about 5% of the eggs when egg-laying sites are limiting, whereas this proportion increased to about 20% when egg-laying sites become abundant. Furthermore, diet manipulation and artificial selection experiments also showed that the exaggerated leg length in M. longipes males is influenced by both genetic and nutritional factors. Collectively, our results highlight how fluctuation in the strength of directional sexual selection, through changes in the intensity of male competition, can drive the exaggeration and phenotypic variation in this weapon trait.