Sexual dimorphism,body size,bite force and male mating success in tuatara

Sexual dimorphisms in body size and head size are common among lizards and are often related to sexual selection on male fighting capacity (organismal performance) and territory defence. However, whether this is generally true or restricted to lizards remains untested. Here we provide data on body and head size, bite performance and indicators of mating success in the tuatara (Sphenodon punctatus), the closest living relative to squamates, to explore the generality of these patterns. First, we test whether male and female tuatara are dimorphic in head dimensions and bite force, independent of body size. Next, we explore which traits best predict bite force capacity in males and females. Finally, we test whether male bite force is correlated with male mating success in a free‐ranging population of tuatara (Sphenodon punctatus). Our data confirm that tuatara are indeed dimorphic in head shape, with males having bigger heads and higher bite forces than females. Across all individuals, head length and the jaw closing in‐lever are the best predictors of bite force. In addition, our data show that males that are mated have higher absolute but not relative bite forces. Bite force was also significantly correlated to condition in males but not females. Whereas these data suggest that bite force may be under sexual selection in tuatara, they also indicate that body size may be the key trait under selection in contrast to what is observed in squamates that defend territories or resources by biting. © 2010 The Linnean Society of London, Biological Journal of the Linnean Society, 2010, 100 , 287–292.     

The Payoff of Fighting in House Crickets: Motivational Asymmetry Increases Male Aggression and Mating Success

Motivational asymmetry caused by differences in subjective resource value is a key component of strategic models of aggression. We study the role of motivational asymmetry in determining differential aggression and mating success of male house crickets, Acheta domesticus. We also assess the extent to which mating differences associated with motivational asymmetry are due to direct male–male fighting vs. male–female interactions. We manipulated male motivation to compete for a mating opportunity by providing males with either no access or nightly access to females for 4 d prior to the experiment. As predicted, when males from each treatment had to compete for the female, those with lower prior access were more aggressive and mated more often. In contrast, when males from each treatment were paired individually with females, there was no significant difference in the frequencies with which they pursued, courted or mated with females. We also found no evidence for female choice based on motivational asymmetry; the rate of successful courtship did not differ between treatments. We conclude that prior mate encounter rate can generate motivational asymmetry, leading to differential mating success mediated by direct male–male aggression.     

Differential aging of bite and jump performance in virgin and mated Teleogryllus commodus crickets

Evolutionary theories of aging state that the force of natural selection declines with age, resulting in trait senescence. However, sexual selection theory predicts that costly traits that signal mate value should increase in expression as survival prospects decline. Mortality rates and fertility tend to show strong signatures of senescence, whereas sexual signaling traits increase with age, but how the expression of traits such as whole-organism performance measures that are subject to both sexual and nonsexual selection should change with age is unclear. We examined the effects of both a key life-history event (mating) and diet quality (male and female optimal diets) on aging in two whole-organism performance traits (bite force and jump take-off velocity) in male and female Teleogryllus commodus crickets. We found no evidence for diet effects on any of the measured traits. Aging effects were more evident in females than in males for both jumping and biting, and constitute a mix of senescence and terminal investment patterns depending on sex/mating class. Sex and mating therefore have important implications for resource allocation to performance traits, and hence for aging of those traits, and interactions between these two factors can result in complex changes in trait expression over individual lifetimes.     

Sexual dimorphism in conspicuousness and ornamentation in the enigmatic leaf‐nosed lizard Ceratophora tennentii from Sri Lanka

Measures of physiological performance capacity, such as bite force, form the functional basis of sexual selection. Information about fighting ability may be conveyed through a structural feature such as a rostrum (i.e. horn) or a colour signal and thereby help reduce costly conflict. We quantified sexual dimorphism in key traits likely to be the targets of sexual selection in Tennent's leaf‐nosed lizard (Ceratophora tennentii) from Sri Lanka, and examined their relationship to bite force and body condition. We found body length and bite force to be similar for males and females. However, head length was significantly greater in males and they had significantly more conspicuous throats and labials (chromatic contrast and luminance) than females. Males also had a proportionally larger rostrum, which we predicted could be an important source of information about male quality for both sexes. Rostrum length was correlated with throat chromatic contrast in males but not females. Nonetheless, the rostrum and aspects of coloration did not correlate with bite force or body condition as we predicted. We have no information on contest escalation in this species but if they rarely bite, as suggested by a lack of difference in bite force between males and females, then bite force and any associated signals would not be a target of selection. Finally, males and females had similar spectral reflectance of the mouth and tongue and both had a peak in the ultra‐violet, and were conspicuous to birds. Lizards only gaped their mouths during capture and not when threatened by a potential predator (hand waving). We hypothesize that conspicuous mouth colour may act as a deimatic signal, startling a potential predator, although this will need careful experimental testing in the future.     

Age at mating and male quality influence female patterns of reproductive investment and survival

The trade‐off between the allocation of resources toward somatic maintenance or reproduction is one of the fundamentals of life history theory and predicts that females invest in offspring at the expense of their longevity or vice versa. Mate quality may also affect life history trade‐offs through mechanisms of sexual conflict; however, few studies have examined the interaction between mate quality and age at first mating in reproductive decisions. Using house crickets (Acheta domesticus), this study examines how survival and reproductive trade‐offs change based on females’ age at first reproduction and exposure to males of varying size. Females were exposed to either a large (presumably high‐quality) or small male at an early (young), middle (intermediate), or advanced (old) age, and longevity and reproductive investment were subsequently tracked. Females mated at a young age had the largest number of eggs but the shortest total lifespans while females mated at older ages produced fewer eggs but had longer total lifespans. The trade‐off between age at first mating and eggs laid appears to be mediated through higher egg‐laying rates and shorter postmating lifespans in females mated later in life. Exposure to small males resulted in shorter lifespans and higher egg‐laying rates for all females indicating that male manipulation of females, presumably through spermatophore contents, varies with male size in this species. Together, these data strongly support a trade‐off between age at first reproduction and lifespan and support the role of sexual conflict in shaping patterns of reproduction.     

Courtship and copulation, but not ejaculates, reduce the longevity of female field crickets (Gryllus bimaculatus)

Females of many taxa incur fitness costs from male sexual coercion and harassment leading to mating. Although male crickets cannot force copulations on females, female Gryllus bimaculatus in this study incurred significant reductions in longevity through being exposed to different levels of male courtship. Virgin females kept in isolation had the longest life spans. Reductions in longevity applied to females in sensory contact with males (without the opportunity to mate), females that courted and mated and females that mated but with fertilization being prevented. Females also incurred significant reductions in longevity when kept with other females, which may have been due to high levels of cannibalism. Consistent with previous studies, females appeared to incur no cost to longevity from receiving sperm or seminal fluid. It is known that female G. bimaculatus benefit genetically from multiple mating. However, this benefit could possibly be offset by the negative effect that male courtship and mating behaviour has on female longevity.     

Sexual selection in hermit crabs: a review and outlines of future research

The information currently available on sexual selection in hermit crabs is reviewed to identify the role of males and females before, during and after mating. According to this information, possible mechanisms of male–male competition, female choice and/or sexual conflict are suggested. Important male components that may affect mating success include dragging the female shell, rotations of the female's shell and male cheliped palpations, and male size and/or shell characteristics (species and size). Possible female determinants of male mating/fertilization success include size (as an indicator of egg production capacity), signalling of sexual receptivity to males, delay from mate guarding to copulation and mating duration. Avenues for deeper exploration in males include the role of the number and morphometry of male sexual tubes during sperm transfer, and whether ejaculate size and sperm number can be adjusted with variable situations of sperm competition intensity and risk. In females it would be interesting to investigate the chemical and behavioural mechanisms affecting spermatophore breakage for sperm release and the variable duration from sperm transfer to spawning. Given these possibilities, and that sperm is externally deposited on the female's body but inside her shell (except for those species that do not use shells, e.g. Birgus , or species where shells are rather small and do not cover the body totally, e.g. Parapagurus ), we conclude that hermit crabs are unique subjects for separating male and female effects, particularly with respect to the applicability of current ideas in sexual selection such as female choice and sexual conflict. Some practical ideas are provided to disentangle both hypotheses using these animals.     

Reversed sexual conflict in a promiscuous antelope

A general tenet of sexual conflict theory is that males have higher optimum mating rates than do females and therefore should be more persistent when it comes to mating. However, in promiscuous species, females might benefit from high mating rates as a result of increased conception probability with favored males, whereas favored males benefit from mating selectively because of sperm depletion. When this results in higher optimum mating rates for females than for males, there is potential for reversed sexual conflicts between persistent females and resistant males. Here I report evidence of such a reversed sexual conflict in a promiscuous antelope, the African topi. Rather than mating randomly, favored males prefer to balance mating investment equally between females as predicted by strategic sperm allocation theory. Females, however, enhance their probability of mating with favored males through aggression toward mating pairs. Supporting the idea that aggressive females thereby harass males to mate at a rate that is suboptimal from the males' perspective, males become increasingly likely to counterattack aggressive females with whom they have already mated disproportionately, and such male counterattacks are associated with refusal to mate with the aggressive females. This study points to reversed sexual conflict as a more significant evolutionary force in promiscuous mammals than previously thought; however, such conflicts probably often go unnoticed because males, in contrast to females, can avoid mating without conspicuous resistance.     

A novel method of comparing mating success and survival reveals similar sexual and viability selection for mobility traits in female tree crickets

The relationship between sexual and viability selection in females is necessarily different than that in males, as investment in sexual traits potentially comes at the expense of both fecundity and survival. Accordingly, females do not usually invest in sexually selected traits. However, direct benefits obtained from mating, such as nuptial gifts, may encourage competition among females and subsidize investment into sexually selected traits. We compared sexual and viability selection on female tree crickets Oecanthus nigricornis, a species where females mate frequently to obtain nuptial gifts and sexual selection on females is likely. If male choice determines female mating success in this species, we expect sexual selection for fecundity traits, as males of many species prefer more fecund females. Alternatively, intrasexual scramble or combat competition on females may select for larger jumping legs or wider heads (respectively). We estimated mating success in wild caught crickets using microsatellite analysis of stored sperm and estimated relative viability by comparing surviving female O. nigricornis to those captured by a common wasp predator. In support of the scramble competition hypothesis, we found sexual selection for females with larger hind legs and narrower heads. We also found stabilizing viability selection for intermediate head width and hind leg size. As predicted, traits under viability and sexual selection were very similar, and the direction of that selection was not opposing. However, because the shape of sexual and viability selection differs, these episodes of selection may favour slightly different trait sizes.     

Seasonal Variation in Female Response to Male Calling Song in the Field Cricket, Gryllus rubens

Conspicuous traits that make males attractive to females may make them vulnerable to predators. Females that approach conspicuous males may increase their risk of predation. This means that selection for reduced male conspicuousness in the presence of predators may be due to sexual selection resulting from altered female behavior in the face of increased predator risk. We examine this hypothesis in the field cricket, Gryllus rubens, in which male calling song attracts both conspecific females for mating and parasitoid flies (Ormia ochracea) which kill their hosts within a week. Female crickets are also parasitized by these flies as a result of associating with calling males. In northern Florida crickets that emerge in the spring are not subject to fly parasitism whereas autumn crickets encounter large numbers of flies. We predicted that autumn females should be less attracted to male song than spring females. We tested female response to male calls in a rectangular arena in which male calling song was broadcast from a speaker. Spring females readily approached the speaker but autumn females were less likely to approach and remain in the vicinity of the speaker. These results emphasize the importance of considering how risk affects the evolution of conspicuous male behavior both directly through its effect on the male and indirectly through its effect on female responses to males.     

Toward a New Sexual Selection Paradigm: Polyandry, Conflict and Incompatibility (Invited Article)

Darwin's recognition that male–male competition and female choice could favor the evolution of exaggerated male traits detrimental to survival set the stage for more than a century of theoretical and empirical work on sexual selection. While this Darwinian paradigm represents one of the most profound insights in biology, its preoccupation with sexual selection as a directional evolutionary force acting on males has diverted attention away from the selective processes acting on females. Our understanding of female reproduction has been further confounded by discreet female mating tactics that have perpetuated the illusion of the monogamous female and masked the potential for conflict between the sexes. With advances in molecular techniques leading to the discovery that polyandry is a pervasive mating strategy, recognition of these shortcomings has brought the study of sexual selection to its current state of flux. In this paper, we suggest that progress in two key areas is critical to formulation of a more inclusive, sexual selection paradigm that adequately incorporates selection from the female perspective. First, we need to develop a better understanding of male × female and maternal × paternal genome interactions and the role that polyandry plays in providing females with non‐additive genetic benefits such as incompatibility avoidance. Consideration of these interaction effects influencing natural selection on females is important because they can complicate and even undermine directional sexual selection on males. Secondly, because antagonistic coevolution maintains a balance between opposing sides that obscures the conflict itself, many more experimental evolution studies and interventionist investigations (e.g. gene knockouts) are needed to tease apart male manipulative adaptations and female counter‐adaptations. It seems evident that the divisiveness and controversy that has plagued sexual selection theory since Darwin first proposed the idea has often stalled progress in this important field of evolutionary biology. What is now needed is a more pluralistic and integrative approach that considers natural as well as sexual selection acting on females, incorporates multiple sexual selection mechanisms, and exploits advances in physiology and molecular biology to understand the mechanisms through which males and females achieve reproductive success.     

Conflict over multiple-partner mating between males and females of the polygynandrous common lizards

The optimal number of mate partners for females rarely coincides with that for males, leading to a potential sexual conflict over multiple-partner mating. This suggests that the population sex ratio may affect multiple-partner mating and thus multiple paternity. We investigate the relationship between multiple paternity and the population sex ratio in the polygynandrous common lizard (Lacerta vivipara). In six populations the adult sex ratio was biased toward males, and in another six populations the adult sex ratio was biased toward females, the latter corresponding to the average adult sex ratio encountered in natural populations. In males the frequency and the degree of polygyny were lower in male-biased populations, as expected if competition among males determines polygyny. In females the frequency of polyandry was not different between treatments, and polyandrous females produced larger clutches, suggesting that polyandry might be adaptive. However, in male-biased populations females suffered from reduced reproductive success compared to female-biased populations, and the number of mate partners increased with female body size in polyandrous females. Polyandrous females of male-biased populations showed disproportionately more mating scars, indicating that polyandrous females of male-biased populations had more interactions with males and suggesting that the degree of multiple paternity is controlled by male sexual harassment. Our results thus imply that polyandry may be hierarchically controlled, with females controlling when to mate with multiple partners and male sexual harassment being a proximate determinant of the degree of multiple paternity. The results are also consistent with a sexual conflict in which male behaviors are harmful to females.     

Increased opportunity for sexual conflict promotes harmful males with elevated courtship frequencies

Mating systems have a profound influence on the probability of conflict occurring between the sexes. Promiscuity is predicted to generate sexual conflict, thereby driving the evolution of male traits that harm females, whereas monogamy is expected to foster reproductive cooperation, thus rendering such traits redundant. We tested these predictions using experimentally evolved Drosophila pseudoobscura subject to different mating systems. Female survival was not influenced by the mating system treatment of her partner. However, females continuously housed with males evolving under elevated opportunities for female promiscuity produced fewer total progeny, but a relatively greater number of progeny early in their lives, than females housed with males evolving under obligate monogamy. We also found that promiscuous males courted females more frequently than monogamous males. Variation in male courtship frequency and progeny production patterns among treatments reinforces the critical importance of mating system variation for sexual conflict, during both pre‐ and post‐copulatory interactions.     

Sexual conflict and ecology: Species composition and male density interact to reduce male mating harassment and increase female survival

Sexual conflict is a pervasive evolutionary force that can reduce female fitness. Experimental evolution studies in the laboratory might overestimate the importance of sexual conflict because the ecological conditions in such settings typically include only a single species. Here, we experimentally manipulated conspecific male density (high or low) and species composition (sympatric or allopatric) to investigate how ecological conditions affect female survival in a sexually dimorphic insect, the banded demoiselle (Calopteryx splendens). Female survival was strongly influenced by an interaction between male density and species composition. Specifically, at low conspecific male density, female survival increased in the presence of heterospecific males (C. virgo). Behavioral mating experiments showed that interspecific interference competition reduced conspecific male mating success with large females. These findings suggest that reproductive interference competition between con‐ and heterospecific males might indirectly facilitate female survival by reducing mating harassment from conspecific males. Hence, interspecific competitors can show contrasting effects on the two sexes thereby influencing sexual conflict dynamics. Our results call for incorporation of more ecological realism in sexual conflict research, particularly how local community context and reproductive interference competition between heterospecific males can affect female fitness.     

The Starling Mating System as an Outcome of the Sexual Conflict

Many bird species demonstrate a variable mating system, with some males being monogamously mated and other males able to attract more than one mate. This variation in avian mating systems is often explained in terms of potential costs of sharing breeding partners and compensation for such costs. However, whenever there is a difference in the optimal mating system for males and females, a sexual conflict over the number of partners is expected. This paper contains a verbal model of how a conflict between male and female European starlings (Sturnus vulgaris),resulting from the fitness consequences of different mating systems for males and females differing over time, determines the mating system. We demonstrate that males and females have contrasting fitness interests regarding mating system, such that males gain from attracting additional mates whereas already mated females pay a cost in terms of reduced reproductive success if males are successful in attracting more mates. We demonstrate how this can be traced to the rules by which males allocate non-sharable care between different broods. Furthermore, we demonstrate that there exist male and female conflict behaviours with the potential to affect the mating system. For example, aggression from already mated females towards prospecting females can limit male mating success and males can circumvent this by spacing the nest-sites they defend. The realised mating system will emerge as a consequence of both the fitness value of the different mating systems for males and females, and the costs for males and females of intersexual competition. We discuss how this model can be developed and critically evaluated in the future.     

Sex peptide causes mating costs in female Drosophila melanogaster

Conflicts between females and males over reproductive decisions are common . In Drosophila, as in many other organisms, there is often a conflict over how often to mate. The mating frequency that maximizes male reproductive success is higher than that which maximizes female reproductive success . In addition, frequent mating reduces female lifespan and reproductive success , a cost that is mediated by male ejaculate accessory gland proteins (Acps) . We demonstrate here that a single Acp, the sex peptide (SP or Acp70A), which decreases female receptivity and stimulates egg production in the first matings of virgin females , is a major contributor to Acp-mediated mating costs in females. Females continuously exposed to SP-deficient males (which produce no detectable SP ) had significantly higher fitness and higher lifetime reproductive success than control females. Hence, rather than benefiting both sexes, receipt of SP decreases female fitness, making SP the first identified gene that is likely to play a central role in sexual conflict.     

Male-male competition and mating success in the orb-web spider,Nephila clavata,with reference to temporal factors

Seasonal occurrence patterns of adults of both sexes, intensity of male-male interactions, and mating success in the spider,Nephila clavata, were examined in the field. Adult males began to attend female webs about 2 weeks before female maturation. Large adult males were abundant in the early breeding season, but small males increased later in the season. From the distribution of males among female webs and size relationship of males within a web, male-male interactions seemed to be more intense when most females were still subadult. This was verified by a field experiment in which males were artificially introduced to female webs that were attended by other males. It was found that the probability of introduced males remaining on subadult female webs was lower than that on adult webs. As mating occurred mostly in the period shortly after the female final molt and first male sperm precedence was known in all spiders reported so far, intense male-male competition on subadult female webs seemed to be reasonable. Male longevity had an important influence on the mating success of males with just-molted females. Mating success was also affected by the relative body size of males present in a given period. Since larger males occupied the position closest to females within a web and stayed there longer, relative body size appeared to influence mating success through male-male competition. Female body size at maturation declined with time; hence, males that attained sexual maturity earlier had the advantage of mating with larger and more fecund females. Therefore, early maturation as well as larger size seem to be two important trairs influencing the reproductive success of males.     

The combined effects of pre‐ and post‐copulatory processes are masking sexual conflict over mating rate in Gerris buenoi

In polygynandrous animals, post‐copulatory processes likely interfere with precopulatory sexual selection. In water striders, sexual conflict over mating rate and post‐copulatory processes are well documented, but their combined effect on reproductive success has seldom been investigated. We combine genetic parentage analyses and behavioural observations conducted in a competitive reproductive environment to investigate how pre‐ and post‐copulatory processes influence reproductive success in Gerris buenoi Kirkaldy. Precopulatory struggles had antagonistic effects on male and female reproductive success: efficiently gaining copulations was beneficial for males, whereas efficiently avoiding copulations was profitable for females. Also, high mating rates and an intermediate optimal resistance level of females supported the hypothesis of convenience polyandry. Contrary to formal predictions, high mating rates (i.e. the number of copulations) did not increase reproductive success in males or decrease reproductive success in females. Instead, the reproductive success of both sexes was higher when offspring were produced with several partners and when there were few unnecessary matings. Thus, male and female G. buenoi displayed different interests in reproduction, but post‐copulatory processes were masking the effects of copulatory mating success on reproductive success. Given the high mating rates observed, sperm competition could easily counter the effect of mating rates, perhaps in interaction with cryptic female choice and/or fecundity selection. Our study presents a complex but realistic overview of sexual selection forces at work in a model organism for the study of sexual conflict, confirming that insights are gained from investigating all episodes in the reproduction cycle of polygynandrous animals.     

Coping with the cold: temperature and mating activity of male sagebrush crickets Cyphoderris strepitans (Orthoptera: Haglidae)

Abstract. The effect of temperature on male sexual displays is well documented but its role in mediating the differential mating success of males is less well established. Male sagebrush crickets Cyphoderris strepitans Morris and Gwynne occur in high‐elevation sagebrush meadows in mountainous areas of the western U.S.A., and often are observed singing at temperatures as low ?8 to ?4 °C to attract sexually receptive females. Males exhibit differential mating success based on their previous mating experience: virgin males have a higher probability of obtaining a mating than do non‐virgin males of securing an additional mating. This differential mating success is mediated in part by an unusual form of nuptial feeding behaviour. Females feed on males’ hind wings at mating and the costs imposed on males as a consequence of this behaviour constrain male mating success. In the present study, to determine whether sagebrush crickets are capable of some degree of thermoregulation, and also to investigate whether this ability is influenced by male mating experience, thoracic temperatures of males of varying mating status were measured in relation to ambient temperature throughout the breeding season. A full factorial model examining the effect of site of capture, mating status, body mass and ambient temperature reveals that only ambient temperature has a significant effect on thoracic temperature after controlling for all other factors. It is concluded that differences in the calling times of virgin and non‐virgin males and their ability to secure mates does not result from differences in their ability to cope physiologically with low temperatures.     

Pre‐ and postcopulatory sexual selection favor aggressive,young males in polyandrous groups of red junglefowl

A challenge in evolutionary biology is to understand the operation of sexual selection on males in polyandrous groups, where sexual selection occurs before and after mating. Here, we combine fine‐grained behavioral information (>41,000 interactions) with molecular parentage data to study sexual selection in replicated, age‐structured groups of polyandrous red junglefowl, Gallus gallus. Male reproductive success was determined by the number of females mated (precopulatory sexual selection) and his paternity share, which was driven by the polyandry of his female partners (postcopulatory sexual selection). Pre‐ and postcopulatory components of male reproductive success covaried positively; males with high mating success also had high paternity share. Two male phenotypes affected male pre‐ and postcopulatory performance: average aggressiveness toward rival males and age. Aggressive males mated with more females and more often with individual females, resulting in higher sexual exclusivity. Similarly, younger males mated with more females and more often with individual females, suffering less intense sperm competition than older males. Older males had a lower paternity share even allowing for their limited sexual exclusivity, indicating they may produce less competitive ejaculates. These results show that—in these populations—postcopulatory sexual selection reinforces precopulatory sexual selection, consistently promoting younger and more aggressive males.