Meek Males and Fighting Females: Sexually-Dimorphic Antipredator Behavior and Locomotor Performance Is Explained by Morphology in Bark Scorpions (Centruroides vittatus)

Sexual dimorphism can result from sexual or ecological selective pressures, but the importance of alternative reproductive roles and trait compensation in generating phenotypic differences between the sexes is poorly understood. We evaluated morphological and behavioral sexual dimorphism in striped bark scorpions (Centruroides vittatus). We propose that reproductive roles have driven sexually dimorphic body mass in this species which produces sex differences in locomotor performance. Poor locomotor performance in the females (due to the burden of being gravid) favors compensatory aggression as part of an alternative defensive strategy, while male morphology is coadapted to support a sprinting-based defensive strategy. We tested the effects of sex and morphology on stinging and sprinting performance and characterized overall differences between the sexes in aggressiveness towards simulated threats. Greater body mass was associated with higher sting rates and slower sprinting within sexes, which explained the greater aggression of females (the heavier sex) and, along with longer legs in males, the improved sprint performance in males. These findings suggest females are aggressive to compensate for locomotor costs of reproduction while males possess longer legs to enhance sprinting for predator evasion and mate finding. Sexual dimorphism in the metasoma (“tail”) was unrelated to stinging and sprinting performance and may best be explained by sexual selection.     

Sexual dimorphism and interpopulation differences in lizard hind limb length: locomotor performance or chemical signalling?

Intraspecific variation in morphology has often been related to fitness differences through its effects on performance. In lizards, variation in hind limb length can be shaped by natural selection for increased locomotor performance, sexual selection on the number or size of femoral pores involved in chemical signalling, or both. Here, we analyse the selective forces involved in sexual dimorphism and differences in hind limb length between two populations of Psammodromus algirus living at different elevation. Males were more robust and had longer hind limbs and limb segments than females, and low‐elevation lizards had longer limbs than high‐elevation lizards. However, differences in locomotor performance were small and non‐significant, making natural selection for faster runs an unlikely explanation for the observed pattern. On the other hand, males had more femoral pores than females, and lizards had more pores at lower elevation, although the difference was significant only for males (which invest more in chemical signalling). In males, the number of pores, which remains constant along a lizard's life, was not correlated with hind limb length. However, femur length was positively correlated with mean pore size, allowing low‐elevation males to have larger than expected pores, which could increase the effectiveness with which they spread their signals in a dry and warm habitat where chemicals become volatile rapidly. Also, saturation of the sexual coloration of the head was higher for low‐elevation males, suggesting that sexual selection pressures may be more intense. Overall, our results indicate that sexual selection plays a significant role in shaping intraspecific variation in hind limb length. © 2011 The Linnean Society of London, Biological Journal of the Linnean Society, 2011, 104 , 318–329.     

It takes two: Seasonal variation in sexually dimorphic weaponry results from divergent changes in males and females

Sexually dimorphic weaponry often results from intrasexual selection, and weapon size can vary seasonally when costs of bearing the weapon exceed the benefits outside of the reproductive season. Weapons can also be favored in competition over nonreproductive resources such as food or shelter, and if such nonreproductive competition occurs year‐round, weapons may be less likely to vary seasonally. In snapping shrimp (Alpheus angulosus), both sexes have an enlarged snapping claw (a potentially deadly weapon), and males of many species have larger claws than females, although females are more aggressive. This contrasting sexual dimorphism (larger weaponry in males, higher aggression in females) raises the question of whether weaponry and aggression are favored by the same mechanisms in males and females. We used field data to determine whether either sex shows seasonal variation in claw size such as described above. We found sexual dimorphism increased during the reproductive season due to opposing changes in both male and female claw size. Males had larger claws during the reproductive season than during the nonreproductive season, a pattern consistent with sexual selection. Females, however, had larger claws during the nonreproductive season than during the reproductive season—a previously unknown pattern of variation in weapon size. The observed changes in female weapon size suggest a trade‐off between claw growth and reproduction in the reproductive season, with investment in claw growth primarily in the nonreproductive season. Sexually dimorphic weaponry in snapping shrimp, then, varies seasonally due to sex differences in seasonal patterns of investment in claw growth, suggesting claws may be advantageous for both sexes but in different contexts. Thus, understanding sexual dimorphisms through the lens of one sex yields an incomplete understanding of the factors favoring their evolution.     

Sexual dimorphism and sexual selection in a montane scincid lizard (Eulamprus leuraensis)

Sex‐based divergences in body sizes and/or shapes within a species imply that selective forces act differently on morphological features in males versus females. That prediction can be tested with data on the relationship between morphology and reproductive output in females, and between morphology and realized paternity (based on genetic assignment tests) in males. In a sample of 81 field‐collected adult Blue Mountains water skinks (Eulamprus leuraensis), males and females averaged similar overall body sizes (snout–vent lengths (SVLs)). Reproductive success (based on 105 progeny produced by the females) increased with SVL at similar rates in both sexes (as expected from the lack of sexual size dimorphism). Multiple paternity was common. Males had larger heads than females of the same body size, and (as predicted) reproductive success increased with relative head size in males but not in females. Males also had relatively longer limbs and shorter trunks than females, but we did not detect significant sex differences in selection on those traits. Reproductive success in both sexes was increased by relatively longer hind limbs. Our data clarify mating systems in this endangered species, and suggest that mating systems are diverse within the genus Eulamprus.     

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.     

Sexual dimorphism in traits related to locomotion: ontogenetic patterns of variation in Podarcis wall lizards

Sexual dimorphism in body size and shape in animals is normally linked to sexual selection mechanisms that modify the morphological properties of each sex. However, sexual dimorphism of ecologically relevant traits may be amplified by natural selection and result in the ecological segregation of both sexes. In the present study, we investigated patterns of sexual dimorphism of morphological traits relevant for locomotion in two lacertid lizards, Podarcis bocagei and Podarcis carbonelli, aiming to identify ontogenetic sources of variation. We analysed trunk and limb variation in relation to total body size, as well as the covariation of different traits, aiming to shed light on the proximate causation of adult sexual dimorphism. We find that, although immatures are generally monomorphic, adult females have a longer trunk, and adult males have longer fore and hind limbs. Both sexes differ substantially with respect to their growth trajectories and relationships between traits, whereas, in some cases, there are signs of morphological constraints delimiting the observed patterns. Because of the direct connection between limb size/shape and locomotor performance, which is relevant both for habitat use and escape from predators, the observed patterns of sexual dimorphism are expected to translate into ecological differences between both sexes. © 2010 The Linnean Society of London, Biological Journal of the Linnean Society, 2010, 99 , 530–543.     

Ecology,sexual dimorphism,and jumping evolution in anurans

Sexual dimorphism (SD) is a common feature of animals, and selection for sexually dimorphic traits may affect both functional morphological traits and organismal performance. Trait evolution through natural selection can also vary across environments. However, whether the evolution of organismal performance is distinct between the sexes is rarely tested in a phylogenetic comparative context. Anurans commonly exhibit sexual size dimorphism, which may affect jumping performance given the effects of body size on locomotion. They also live in a wide variety of microhabitats. Yet the relationships among dimorphism, performance, and ecology remain underexamined in anurans. Here, we explore relationships between microhabitat use, body size, and jumping performance in males and females to determine the drivers of dimorphic patterns in jumping performance. Using methods for predicting jumping performance through anatomical measurements, we describe how fecundity selection and natural selection associated with body size and microhabitat have likely shaped female jumping performance. We found that the magnitude of sexual size dimorphism (where females are about 14% larger than males) was much lower than dimorphism in muscle volume, where females had 42% more muscle than males (after accounting for body size). Despite these sometimes-large averages, phylogenetic t-tests failed to show the statistical significance of SD for any variable, indicating sexually dimorphic species tend to be closely related. While SD of jumping performance did not vary among microhabitats, we found female jumping velocity and energy differed across microhabitats. Overall, our findings indicate that differences in sex-specific reproductive roles, size, jumping-related morphology, and performance are all important determinants in how selection has led to the incredible ecophenotypic diversity of anurans.     

Sex-specific selective pressures on body mass in the greater white-toothed shrew, Crocidura russula

The direction, intensity and shape of viability-, sexual- and fecundity selection on body mass were investigated in a natural population of the greater white-toothed shrew (Crocidura russula), combining parentage assignment through molecular techniques and mark-recapture data over several generations. A highly significant stabilizing viability selection was found in both sexes, presumably stemming from the constraints imposed by their insectivorous habits and high metabolic costs. Sexual selection, directional in both sexes, was twice as large in males than in females. Our results suggest that body mass matters in this context by facilitating the acquisition and defense of a breeding territory. No fecundity selection could be detected. The direction of sexual size dimorphism (SSD) was in agreement with the observed pattern of selective pressures: males were heavier than females, because of stronger sexual selection. SSD intensity, however, was low compared with other mammals, because of the low level of polygyny, the active role of females in territory defense and the intensity of stabilizing viability selection.     

MALE‐BIASED FITNESS EFFECTS OF SPONTANEOUS MUTATIONS IN DROSOPHILA MELANOGASTER

In populations with males and females, sexual selection may often represent a major component of overall selection. Sexual selection could act to eliminate deleterious alleles in concert with other forms of selection, thereby improving the fitness of sexual populations. Alternatively, the divergent reproductive strategies of the sexes could promote the maintenance of sexually antagonistic variation, causing sexual populations to be less fit. The net impact of sexual selection on fitness is not well understood, due in part to limited data on the sex‐specific effects of spontaneous mutations on total fitness. Using a set of mutation accumulation lines of Drosophila melanogaster, we found that mutations were deleterious in both sexes and had larger effects on fitness in males than in females. This pattern is expected to reduce the mutation load of sexual females and promote the maintenance of sexual reproduction.     

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.     

SEXUAL SELECTION AFFECTS THE EVOLUTION OF LIFESPAN AND AGEING IN THE DECORATED CRICKET GRYLLODES SIGILLATUS

Recent work suggests that sexual selection can influence the evolution of ageing and lifespan by shaping the optimal timing and relative costliness of reproductive effort in the sexes. We used inbred lines of the decorated cricket, Gryllodes sigillatus, to estimate the genetic (co)variance between age‐dependent reproductive effort, lifespan, and ageing within and between the sexes. Sexual selection theory predicts that males should die sooner and age more rapidly than females. However, a reversal of this pattern may be favored if reproductive effort increases with age in males but not in females. We found that male calling effort increased with age, whereas female fecundity decreased, and that males lived longer and aged more slowly than females. These divergent life‐history strategies were underpinned by a positive genetic correlation between early‐life reproductive effort and ageing rate in both sexes, although this relationship was stronger in females. Despite these sex differences in life‐history schedules, age‐dependent reproductive effort, lifespan, and ageing exhibited strong positive intersexual genetic correlations. This should, in theory, constrain the independent evolution of these traits in the sexes and may promote intralocus sexual conflict. Our study highlights the importance of sexual selection to the evolution of sex differences in ageing and lifespan in G. sigillatus.     

The postnatal ontogeny of the sexually dimorphic vocal apparatus in goitred gazelles (Gazella subgutturosa)

This study quantitatively documents the progressive development of sexual dimorphism of the vocal organs along the ontogeny of the goitred gazelle (Gazella subgutturosa). The major, male‐specific secondary sexual features, of vocal anatomy in goitred gazelle are an enlarged larynx and a marked laryngeal descent. These features appear to have evolved by sexual selection and may serve as a model for similar events in male humans. Sexual dimorphism of larynx size and larynx position in adult goitred gazelles is more pronounced than in humans, whereas the vocal anatomy of neonate goitred gazelles does not differ between sexes. This study examines the vocal anatomy of 19 (11 male, 8 female) goitred gazelle specimens across three age‐classes, that is, neonates, subadults and mature adults. The postnatal ontogenetic development of the vocal organs up to their respective end states takes considerably longer in males than in females. Both sexes share the same features of vocal morphology but differences emerge in the course of ontogeny, ultimately resulting in the pronounced sexual dimorphism of the vocal apparatus in adults. The main differences comprise larynx size, vocal fold length, vocal tract length, and mobility of the larynx. The resilience of the thyrohyoid ligament and the pharynx, including the soft palate, and the length changes during contraction and relaxation of the extrinsic laryngeal muscles play a decisive role in the mobility of the larynx in both sexes but to substantially different degrees in adult females and males. Goitred gazelles are born with an undescended larynx and, therefore, larynx descent has to develop in the course of ontogeny. This might result from a trade‐off between natural selection and sexual selection requiring a temporal separation of different laryngeal functions at birth and shortly after from those later in life. J. Morphol. 277:826–844, 2016. © 2016 Wiley Periodicals, Inc.     

Sexual size dimorphism and selection in the wild in the waterstrider Aquarius remigis: Body size,components of body size and male mating success

Sexual size dimorphism is assumed to be adaptive and is expected to evolve in response to a difference in the net selection pressures on the sexes. Although a demonstration of sexual selection is neither necessary nor sufficient to explain the evolution of sexual size dimorphism, sexual selection is generally assumed to be a major evolutionary force. If contemporary sexual selection is important in the evolution and maintenance of sexual size dimorphism then we expect to see concordance between patterns of sexual selection and patterns of sexual dimorphism. We examined sexual selection in the wild, acting on male body size, and components of body size, in the waterstrider Aquarius remigis, as part of a long term study examining net selection pressures on the two sexes in this species. Selection was estimated on both a daily and annual basis. Since our measure of fitness (mating success) was behavioral, we estimated reliabilities to determine if males perform consistently. Reliabilities were measured as ? statistics and range from fair to perfect agreement with substantial agreement overall. We found significant univariate sexual selection favoring larger total length in the first year of our study but not in the second. Multivariate analysis of components of body size revealed that sexual selection for larger males was not acting directly on total length but on genital length. Sexual selection for larger male body size was opposed by direct selection favoring smaller midfemoral lengths. While males of this species are smaller than females, they have longer genital segments and wider forefemora. Patterns of contemporary sexual selection and sexual size dimorphism agree only for genital length. For total length, and all other components of body size examined, contemporary sexual selection was either nonsignificant or opposed the pattern of size dimporhism. Thus, while the net pressures of contemporary selection for the species may still act to maintain sexual size dimorphism, sexual selection alone does not.     

Sexual size dimorphism is not associated with the evolution of parental care in frogs

Sex differences in parental care are thought to arise from differential selection on the sexes. Sexual dimorphism, including sexual size dimorphism (SSD), is often used as a proxy for sexual selection on males. Some studies have found an association between male‐biased SSD (i.e., males larger than females) and the loss of paternal care. While the relationship between sexual selection on males and parental care evolution has been studied extensively, the relationship between female‐biased SSD (i.e., females larger than males) and the evolution of parental care has received very little attention. Thus, we have little knowledge of whether female‐biased SSD coevolves with parental care. In species displaying female‐biased SSD, we might expect dimorphism to be associated with the evolution of paternal care or perhaps the loss of maternal care. Here, drawing on data for 99 extant frog species, we use comparative methods to evaluate how parental care and female‐biased SSD have evolved over time. Generally, we find no significant correlation between the evolution of parental care and female‐biased SSD in frogs. This suggests that differential selection on body size between the sexes is unlikely to have driven the evolution of parental care in these clades and questions whether we should expect sexual dimorphism to exhibit a general relationship with the evolution of sex differences in parental care.     

Consequences of sexual size dimorphism on energetics and locomotor performance of Grammostola rosea (Araneae; Teraphosidae)

Most male spiders are smaller than females; during sexual maturity, males change their behaviour, abandoning their web or nest to seek out receptive females actively, whereas females stalk prey near their web or nest and tend not to move away from it. Considering this behavioural difference to be associated with increased locomotor activity at maturity, it may be hypothesized that males will have traits that increase locomotor performance. The present study examines the kinetics and energetics of the movements of the mygalomorph spider Grammostola rosea Walckenaer, a large spider with sexual size dimorphism. It is found that males have a higher maximum aerobic speed, average speed, distance travelled and critical angle of climbing than females, indicating better performance. Males also have lower costs of transport than females. These results support the hypothesis that sexual dimorphism in wandering spiders with active males, which are characterized by smaller body size and longer legs than the larger and more static females, is associated with low transport cost, high velocity and better locomotor performance.     

Intrapopulation variation in predator-avoidance performance of Galápagos lava lizards: The interaction of sexual and natural selection

Summary The interaction of sexual and natural selection in shaping variation in defensive behavior was explored via three steps. (1) Three predictions from the hypothesis that varying responses to predation account for intrapopulation variation in locomotory performance and wariness were tested. One measure of locomotory performance and two measures of wariness were compared between lava lizards (Tropidurus albemarlensis) inhabiting sparsely (1% cover) and heavily (33% cover) vegetated areas of Isla Plaza Sur in the Galapagos Archipelago. (2) Variation in morphology was examined to identify proximate mechanisms for differences seen in (1). (3) Levels of predation were compared between males and females to test a prediction from a model explaining the observed variation in defensive behavior. Male, but not female, lizards from sparsely vegetated areas were faster than those from heavily vegetated areas. Both male and female lizards from sparsely vegetated areas were significantly warier than those from heavily vegetated areas. Multiple regression, covariance, and residual analyses identify relatively longer hindlimbs of males as the proximate cause of their greater speed over that of females, rather than differences in body size, but neither body nor hindlimb size account for the microgeographic differences in speed of males. Significantly higher predation on males provided a positive test of the major prediction from our model of selective forces in which sexual selection for territory defense by males favors short approach distances (by minimizing time away from the territory caused by erroneous flight) and leads to natural selection for their increased hindlimb size and speed.     

Sexual Selection of Zorion guttigerum Westwood (Coleoptera: Cerambycidae: Cerambycinae) in Relation to Body Size and Color

Zorion guttigerum is a flower-visiting longhorned beetle endemic to New Zealand. Sexual selection of this species in relation to the body size and color form of different sexes was investigated in the field. The population sex ratio, based on censuses of feeding and mating sites (flowers), is male-biased. Females are significantly larger than males. Both sexes have antennae of similar length but the antennal length relative to the elytral length is greater in males than in females, and the antennal length of males increases more with an increase in body size than that of females. Both sexes have dark blue (DB) and yellowish-brown (YB) individuals. Both pair-bonded and solitary males are similar in elytral and antennal length. In pair-bonded males, DB individuals are significantly more numerous than YB ones, but in solitary males, the number of both color forms is similar. Males tend to have territory protection behavior, fighting with and chasing away rival males from feeding and mating sites. Larger males usually win the fight but the size-dependent fighting advantage does not translate into mating success. Male color plays an important role in mating success, with DB males having a significantly better chance to mate than YB males. Furthermore, male body size and color also have interactions in mating success: males of DB color morph obtain a greater mating advantage according to body size. Pair-bonded females are significantly larger and have longer antennae than solitary females, suggesting that males prefer larger females for mating. In addition, females of DB color morph with longer antennae are also preferred by males for mating. The significance of these findings is discussed.     

Sex‐specific selection and sexual size dimorphism in the waterstrider Aquarius remigis

We estimated selection on adult body size for two generations in two populations of Aquarius remigis, as part of a long‐term study of the adaptive significance of sexual size dimorphism (SSD). Net adult fitness was estimated from the following components: prereproductive survival, daily reproductive success (mating frequency or fecundity), and reproductive lifespan. Standardized selection gradients were estimated for total length and for thorax, abdomen, genital and mesofemur lengths. Although selection was generally weak and showed significant temporal and spatial heterogeneity, patterns were consistent with SSD. Prereproductive survival was strongly influenced by date of eclosion, but size (thorax and genital lengths in females; total and abdomen lengths in males) played a significant secondary role. Sexual selection favoured smaller males with longer external genitalia in one population. Net adult fitness was not significantly related to body size in females, but was negatively related to size (thorax and total length) in males.     

Environmental complexity and the purging of deleterious alleles

Sexual interactions among adults can generate selection on both males and females with genome‐wide consequences. Sexual selection through males is one component of this selection that has been argued to play an important role in purging deleterious alleles. A common technique to assess the influence of sexual selection is by a comparison of experimental evolution under enforced monogamy versus polygamy. Mixed results from past studies may be due to the use of highly simplified laboratory conditions that alter the nature of sexual interactions. Here, we examine the rate of purging of 22 gene disruption mutations in experimental polygamous populations of Drosophila melanogaster in each of two mating environments: a simple, high‐density environment (i.e., typical fly vials), and a lower density, more spatially complex environment. Based on past work, we expect sexual interactions in the latter environment to result in stronger selection in both sexes. Consistent with this, we find that mutations tend to be purged more quickly in populations evolving in complex environments. We discuss possible mechanisms by which environmental complexity might modulate the rate at which deleterious alleles are purged and putatively ascribe a role for sexual interactions in explaining the treatment differences in our experiment.     

Stronger condition dependence in female size explains altitudinal variation in sexual size dimorphism of a Tibetan frog

The present study investigated altitudinal variation in sexual size dimorphism of a Tibetan frog Nanorana parkeri. Size dimorphism was female‐biased in all populations, although this bias became less at higher altitudes because of a steeper altitudinal decrease in female size than male size. Operational sex ratios, an indicator of the opportunity for sexual selection on larger males, changed independently of altitude. Clutch volume, an indicator of the strength of fecundity selection on larger females, was positively with female size, and tended to decrease approaching high altitudes. Females lived longer and grew more slowly than males, and the mean age in both sexes increased and growth rate decreased altitudinally, although the changes were more rapid in females than males. These results suggest that, relative to males, females (i.e. the sex that typically bears greater reproductive costs and experiences stronger directional selection for larger size to take fecundity advantages) should be more sensitive to environments, attaining a larger size via enhancing growth under favourable lower‐latitude conditions but a smaller size as a result of retarding growth when conditions become harsher at higher altitudes. This supports the condition‐dependence hypothesis with respect to intraspecific variation in sexual size dimorphism. © 2012 The Linnean Society of London, Biological Journal of the Linnean Society, 2012, 107 , 558–565.