Exploring sexual differences in external morphology and limb muscles of Hylarana guentheri (Anura: Ranidae) during non-breeding season

Studying sexual dimorphism facilitates better understanding of the general intersexual divergence of the same species and gives insights into the impact of selective forces on each sex. The sexual dimorphism in anuran external morphology or limb muscles has been well studied in reproductive context, but less so outside the breeding season. Herein, the intersexual differences in 12 external morphological characteristics, 11 forelimb and 12 hindlimb muscles of Hylarana guentheri during non-breeding season were investigated. We found that the males possessed significantly greater head width and tympanum diameter, while the females had significantly larger body size and head length; there was no sexual differences in the examined limb muscles, except for the mass of flexor carpi radialis. The larger body size means greater fertility for the females, and the longer head may be related to the allometry and reduce resource competition. For the males, the wider head is likely correlated with prey size and male–male competition, and also, the larger tympanum and heavier flexor carpi radialis probably aid the productive success. This study provides the comprehensive morphological accounts about the sexual differences of H. guentheri during non-breeding season, which will contribute to clarify the sex-specific resource allocation and reproductive strategies of anurans.     

黑眶蟾蜍的两性异形与选型配对模式

本研究以黑眶蟾蜍(Duttaphrynus melanostictus)为研究对象,通过对比黑眶蟾蜍抱对个体的体长、头长、头宽、眼间距、鼓膜径、耳后腺长、眼径、前臂及手长、前肢长以及后肢长等形态特征,分析雌性黑眶蟾蜍繁殖输出与其体型的关系,探究黑眶蟾蜍两性异形模式及其与雌性生育力的关系;同时通过对配对个体形态学特征的相关性分析探究了黑眶蟾蜍的配对模式。结果表明,黑眶蟾蜍雌性体长和体重显著大于雄体;两性的所有局部形态特征均与体长成正相关;去除体长因素影响后,雄性头长以及后肢长均明显大于雌性,其余局部形态特征两性间则皆无显著差异。雌体的窝卵重、窝卵数均与其体长和体重成正相关关系。雌性成体的前肢长与抱对雄性成体的前肢长之间呈显著正相关,其余形态特征两性间均无相关性。研究表明,生育力选择是导致黑眶蟾蜍两性异形的重要驱动力;黑眶蟾蜍的选型配对模式未表现在个体大小上,而是体现在局部特征(前肢长),这不仅为揭示两栖类配对模式的普遍性提供了参考,还表明对两栖类选型配对的研究应以多个性状为对象。     

The short legs of great apes: evidence for aggressive behavior in australopiths

Early hominins, australopiths, were similar to most large primates in having relatively short hindlimbs for their body size. The short legs of large primates are thought to represent specialization for vertical climbing and quadrupedal stability on branches. Although this may be true, there are reasons to suspect that the evolution of short legs may also represent specialization for physical aggression. Fighting in apes is a behavior in which short legs are expected to improve performance by lowering the center of mass during bipedal stance and by increasing the leverage through which muscle forces can be applied to the ground. Among anthropoid primates, body size sexual dimorphism (SSD) and canine height sexual dimorphism (CSD) are strongly correlated with levels of male-male competition, allowing SSD and CSD to be used as indices of male-male aggression. Here I show that the evolution of hindlimb length in apes is inversely correlated with the evolution of SSD (R(2)= 0.683, P-value = 0.006) and the evolution of CSD (R(2)= 0.630, P-value = 0.013). In contrast, a significant correlation was not observed for the relationship between the evolution of hindlimb and forelimb lengths. These observations are consistent with the suggestion that selection for fighting performance has maintained relatively short hindlimbs in species of Hominoidea with high levels of male-male competition. Although australopiths were highly derived for striding bipedalism when traveling on the ground, they retained short legs compared to those of Homo for over two million years, approximately 100,000 generations. Their short legs may be indicative of persistent selection for high levels of aggression.     

Sexual selection on forelimb muscles of western grey kangaroos (Skippy was clearly a female)

Studies of sexual selection have tended to concentrate on obvious morphological dimorphisms such as crests, horns, antlers, and other physical displays or weapons; however, traits that show no obvious sexual dimorphism may nevertheless still be under sexual selection. Sexual selection theory generally predicts positive allometry for sexually selected traits. When fighting, male kangaroos use their forelimbs to clasp and hold their opponent and, standing on their tail, bring up their hind legs to kick their opponent. This action requires substantial strength and balance. We examined allometry of forelimb musculature in male and female western grey kangaroos (Macropus fuliginosus) to determine whether selection through male–male competition is associated with sex differences in muscle development. Forelimbs of males are more exaggerated than in females, with relatively greater muscle mass in males than the equivalent muscles in females. Furthermore, while muscles generally showed isometric growth in female forelimbs, every muscle demonstrated positive allometry in males. The significant positive allometry in male forelimb musculature, particularly those muscles most likely involved in male–male combat (a group of muscles involved in grasping: shoulder adduction, elbow flexion; and pulling: arm retraction, elbow flexion), clearly suggests that this musculature is subject to sexual selection. In addition to contributing to locomotion, the forelimbs of male kangaroos can also act as a signal, a weapon, and help in clasping, features that would contribute towards their importance as a sexually selected trait. Males would therefore benefit from well‐developed musculature of the arms and upper body during competition for mates. © 2013 The Linnean Society of London, Biological Journal of the Linnean Society, 2013, 109 , 923–931.     

Reversed sexual size dimorphism in microtines: Are females larger than males or are males smaller than females?

Summary We analysed sexual size dimorphism for 21 populations of microtine rodents. Female to male size ratio varied considerably among populations from females significantly larger than males (ratio=1.18) to males larger than females (ratio=0.78). In a multiple regression analysis female to male home range size ratio explained 94% of the total variation in body size dimorphism and was the only one of eight independent variables that was selected in a stepwise regression procedure. When females are the larger sex, males have home range sizes much larger than females. We suggest that the relationship between home range size ratio and body weight size dimorphism reflects different selection pressures on males and females in competition for resources and mates.     

Comparison of the effect of castration on the development of postural and non-postural muscles of mice.

The effect of castration on the development of muscle mass of postural and non-postural muscles was studied in 18 male mice (9 castrated, 9 uncastrated). Results obtained indicated that the castrated males grew faster and were bigger in body size and weight at maturity than the intact males. The bigger body size of castrated males was not due to larger muscle mass but was probably due to increased subcutaneous fat deposition. Atrophy of muscles usually observed following castration was significantly greater in the non-postural (biceps brachii) muscle of the forelimb as compared to the postural (triceps brachii) muscle of the forelimb. Conversely, the amount of reduction in muscle mass was similar in both postural (soleus) and non-postural (tibialis cranialis) muscles of the hindlimb.     

Sexual size dimorphism and timing of spring migration in birds

Sexually selected traits are limited by selection against those traits in other fitness components, such as survival. Thus, sexual selection favouring large size in males should be balanced by higher mortality of larger males. However, evidence from red-winged blackbirds (Agelaius phoeniceus) indicates that large males survive better than small males. A survival advantage to large size could result from males migrating north in early spring, when harsh weather favours large size for energetic reasons. From this hypothesis we predicted that, among species, sex differences in body size should be correlated with sex differences in timing of spring migration. The earlier males migrate relative to females, the larger they should be relative to females. We tested this prediction using a comparative analysis of data collected from 30 species of passerine birds captured on migration. After controlling for social mating system, we found that sexual size dimorphism and difference in arrival dates of males and females were significantly positively correlated. This result is consistent with the hypothesis that selection for survival ability promotes sexual size dimorphism (SSD), rather than opposes SSD as is the conventional view. If both natural selection and sexual selection favour large adult males, then limits to male size must be imposed before males become adults.     

Sexual selection, antennae length and the mating advantage of large males in Asellus aquaticus

In crustacean species with precopulatory mate-guarding, sexual size dimorphism has most often been regarded as the consequence of a large male advantage in contest competition for access to females. However, large body size in males may also be favoured indirectly through scramble competition. This might partly be the case if the actual target of selection is a morphological character, closely correlated with body size, involved in the detection of receptive females. We studied sexual selection on body size and antennae length in natural populations of Asellus aquaticus, an isopod species with precopulatory mate guarding. In this species, males are larger than females and male pairing success is positively related to body size. However, males also have longer antennae, relative to body size, than females, suggesting that this character may also be favoured by sexual selection. We used multivariate analysis of selection to assess the relative influences of body size and antennae length in five different populations in the field. Selection gradients indicated that, overall, body size was a better predictor of male pairing success than antennae length, although some variation was observed between sites. We then manipulated male antennae length in a series of experiments conducted in the lab, and compared the pairing ability of males with short or long antennae. Males with short antennae were less likely to detect, orient to, and to pair with a receptive female compared with males with long antennae. We discuss the implications of our results for studies of male body size and sexual dimorphism in relation to sexual selection in crustaceans.     

Sexual selection, antennae length and the mating advantage of large males in Asellus aquaticus

In crustacean species with precopulatory mate-guarding, sexual size dimorphism has most often been regarded as the consequence of a large male advantage in contest competition for access to females. However, large body size in males may also be favoured indirectly through scramble competition. This might partly be the case if the actual target of selection is a morphological character, closely correlated with body size, involved in the detection of receptive females. We studied sexual selection on body size and antennae length in natural populations of Asellus aquaticus, an isopod species with precopulatory mate guarding. In this species, males are larger than females and male pairing success is positively related to body size. However, males also have longer antennae, relative to body size, than females, suggesting that this character may also be favoured by sexual selection. We used multivariate analysis of selection to assess the relative influences of body size and antennae length in five different populations in the field. Selection gradients indicated that overall body size was a better predictor of male pairing success than antennae length, although some variation was observed between sites. We then manipulated male antennae length in a series of experiments conducted in the laboratory, and compared the pairing ability of males with short or long antennae. Males with short antennae were less likely to detect, orient to and to pair with a receptive female compared to males with long antennae. We discuss the implications of our results for studies of male body size and sexual dimorphism in relation to sexual selection in crustaceans.     

Male-biased size dimorphism in ichneumonine wasps (Hymenoptera: Ichneumonidae) – the role of sexual selection for large male size

Abstract.  1. Sexual differences in body size are expected to evolve when selection on female and male sizes favours different optima.
2. Insects have typically female-biased size dimorphism that is usually explained by the strong fecundity advantage of larger size in females. However, numerous exceptions to this general pattern have led to the search for selective pressures favouring larger size in males.
3. In this study, the benefits of large size were investigated in males of four species of ichneumonine wasps, a species-rich group of parasitoids, many representatives of which exhibit male-biased size dimorphism.
4. Mating behaviour of all ichneumonine wasps are characterised by pre-copulatory struggles, in the course of which males attempt to override female reluctance to mate. A series of laboratory trials was conducted to study the determinants of male mating success.
5. A tendency was found for larger males as well as those in better condition to be more successful in achieving copulations. Size dimorphism of the species studied, mostly male-biased in hind tibia length but female-biased in body weight, indicates that sexual selection in males favours longer bodies and appendages rather than larger weight.
6. The qualitative similarity of the mating patterns suggests that sexual selection cannot completely explain the considerable among-species differences in sexual size dimorphism.
7. The present study cautions against using various size indices as equivalents for calculating sexual size dimorphism.
8. It is suggested that female reluctance in ichneumonine wasps functions as a mechanism of female mate assessment.     

Directional changes in sexual size dimorphism in shorebirds, gulls and alcids

The Charadrii (shorebirds, gulls and alcids) are one of the most diverse avian groups from the point of view of sexual size dimorphism, exhibiting extremes in both male-biased and female-biased dimorphism, as well as monomorphism. In this study we use phylogenetic comparative analyses to investigate how size dimorphism has changed over evolutionary time, distinguishing between changes that have occurred in females and in males. Independent contrasts analyses show that both body mass and wing length have been more variable in males than in females. Directional analyses show that male-biased dimorphism has increased after inferred transitions towards more polygynous mating systems. There have been analogous increases in female-biased dimorphism after transitions towards more socially polyandrous mating systems. Changes in dimorphism in both directions are attributable to male body size changing more than female body size. We suggest that this might be because females are under stronger natural selection constraints related to fecundity. Taken together, our results suggest that the observed variation in dimorphism of Charadrii can be best explained by male body size responding more sensitively to variable sexual selection than female body size.     

Sexual behaviour and evolution of sexual dimorphism in body size in Jaera (Isopoda Asellota)

Individuals of the genus Jaera do not mate at random. In the species from the Mediterranean group, J. italica and. J. nordmanni, large males and medium sized females are at an advantage and their sizes are positively assorted. These effects are attributable to sexual competition between males. In the Ponlo-caspian species J. istri, no advantage of large males exists, but sexual selection could be the cause for a long passive phase prior to copulation and for normalizing selection upon female size at pairing. In the Atlantic species, J. albifrons, no selection can be ascertained.
Differential mating success in males appears as one of the causes of the evolution of sexual dimorphism in body size, which makes males larger, of equal size, or smaller than females according to the species. The reason for this reversal in dimorphism seems to differ in the two sexes. Sexual selection provides an explanation for the evolution of male size, while the interspecific changes in female length are more likely due to ecological factors.     

Sexual differences in locomotor performance in Tropidurus catalanensis lizards (Squamata: Tropiduridae) – body shape,size and limb musculature explain variation between males and females

Sexual dimorphism (SD) is the evolutionary outcome of selection acting differently on males and females. Several studies describe sexual differences in body size, although other morphological traits might be allometric between sexes and imply functional consequences. Here we test whether morphological differences between sexes in size and shape in the lizard Tropidurus catalanensis explain variation in performance of four locomotor traits. Our results show that males are larger than females and also exhibit longer limbs, longer muscles and larger muscle cross‐sectional areas, while females have longer trunks and more sharped anterior claws; males outperform females in all locomotor performances measured. Sexual differences in sprinting and climbing is related with body size, and climbing performance is also explained by limb lengths, by differences in lengths and cross‐sectional areas of specific muscles, and by interlimb distances. Between‐sex differences in exertion are also related to SD, despite associations with sharper posterior claws that are independent of sex. Grasping performance, however, is associated with some muscle and morphological parameters that are not sexually dimorphic. Together our results suggest that morphology might be under sexual selection in T. catalanensis, given that better locomotor performance likely favours male lizards in typical activities of this polygenic species, such as territory defence and female acquisition. Moreover, the longer trunks that characterize females may confer more space to accommodate eggs. On the other hand, territory defence by males probably increases their exposure to predators, resulting in a synergistic effect of sexual and natural selection in the evolution of SD in T. catalanensis.     

Sexual size dimorphism and assortative mating in Carolina Wrens

ABSTRACT.   Sexual size dimorphism (SSD) may be due to sexual and natural selection, but identifying specific mechanisms that generate such dimorphism in a species is difficult. I examined SSD in Carolina Wrens ( Thryothorus ludovicianus ) by examining (1) the degree of SSD in the population and between pairs using five morphometrics, (2) assortative mating patterns based on size and age, and (3) relationships between size and longevity. Analysis revealed that males were significantly larger than females in all body measurements. For example, mass, bill, and wing measurements yielded a canonical variable that permitted separation of the sexes and linear classification functions correctly determined the sex of 95% (238/250) of all wrens measured. No evidence was found to suggest that SSD was related to resource partitioning. However, assortative mating trends based on morphometrics (e.g., wing length), positive associations between longevity and morphometrics (e.g., wing length in females and body size in males), and intense male-male contests for territorial resources year-round provide evidence that sexual selection may contribute to SSD in Carolina Wrens.     

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

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

Sexual dimorphism and synchrony of breeding: variation in polygyny potential among populations in the common brushtail possum, Trichosurus vulpecula

Male-biased sexual dimorphism in both size and body mass isa common characteristic of many polygynous mammals and is oftenattributed to sexual selection favoring large males. The degreeof dimorphism is thought to be related to the potential forsome males to monopolize access to estrous females, which isin turn related to the distribution of receptive females inspace and time. In the present study, we investigated the relationshipbetween the temporal distribution of breeding females and thedegree of mass dimorphism among 11 populations of the commonbrushtail possum from northern Australia. Breeding patternsvaried from complete aseasonality in some populations to a seasonalconcentration of births within 2 or 3 months in others. We predictedthat in populations in which mating opportunities were distributedthroughout the year, dominant males would be able to monopolizeaccess to larger numbers of estrous females than in populationsin which matings were more seasonal, and in such populations,large body size in males would be favored. We found that dimorphismwas related to seasonality of breeding, being greatest in populationswith a more aseasonal pattern. Mean body mass of male possumsalso decreased with increasing population density. Populationdensity may influence the degree of breeding synchrony withinpopulations, particularly in locations with a more seasonalclimate. The present study is the first to demonstrate plasticityin mass dimorphism in response to local variation in the synchronyof breeding in a mammal species.     

Female‐biased size dimorphism in a diapausing caddisfly,Mesophylax aspersus: effect of fecundity and natural and sexual selection

1. The effect of mating success, female fecundity and survival probability associated with intra‐sex variation in body size was studied in Mesophylax aspersus, a caddisfly species with female‐biased sexual size dimorphism, which inhabits temporary streams and aestivates in caves. Adults of this species do not feed and females have to mature eggs during aestivation. 2. Thus, females of larger size should have a fitness advantage because they can harbour more energy reserves that could influence fecundity and probability of survival until reproduction. In contrast, males of smaller size might have competitive advantages over others in mating success. 3. These hypotheses were tested by comparing the sex ratio and body size of individuals captured before and after the aestivation period. The associations between body size and female fecundity, and between mating success and body size of males, were explored under laboratory conditions. 4. During the aestivation period, the sex ratio changed from 1 : 1 to male biased (4 : 1), and a directional selection on body size was detected for females but not for males. Moreover, larger clutches were laid by females of larger size. Finally, differences in mating success between small and large males were not detected. These results suggest that natural selection (i.e. the differential mortality of females associated with body size) together with possible fecundity advantages, are important factors responsible of the sexual size dimorphism of M. aspersus. 5. These results highlight the importance of taking into account mechanisms other than those traditionally used to explain sexual dimorphism. Natural selection acting on sources of variation, such as survival, may be as important as fecundity and sexual selection in driving the evolution of sexual size dimorphism.     

Reproductive behavior of the Japanese treefrog,Rhacophorus arboreus (Anura: Rhacophoridae)

Reproductive behavior of the foam-nesting treefrog,Rhacophorus arboreus, is described. Oviposition was made either by an amplectant pair (a female, and an amplectant male) alone or by an amplectant pair and other males (joiners). The snout-vent length of males and females was negatively correlated with the date of the first appearance in the pond. The period of staying in the pond in males was longer than that in females. Physical body contacts of other males, and encounter calls by ampletant males were frequently observed before oviposition with joiners. Snout-vent length of the female and an amplectant male were positively correlated. Encounter calls by amplectant males were likely to precede movement of females. There was no difference of snout-vent length between amplectant males and joiners. Females moved the hindlimbs more frequently than males during oviposition. Amplectant males made encounter cells more frequently in oviposition with joiners, than oviposition without joiners. The patterns of transitions between acts of females and amplectant males were similar between oviposition with and without joiners. Females more frequently responded to amplectant males than joiners. Both the duration of a bout of the movements of hindlimbs of females and the interval between the bouts were longer in oviposition without joiners than that with joiners. The number of the movements per bout in oviposition without joiners was smaller than that with joiners. Encounter calls by amplectant males during oviposition tended to precede the hindlimb movements of females.     

The interplay between natural and sexual selection in the evolution of sexual size dimorphism in Sceloporus lizards (Squamata: Phrynosomatidae)

Sexual size dimorphism (SSD) evolves because body size is usually related to reproductive success through different pathways in females and males. Female body size is strongly correlated with fecundity, while in males, body size is correlated with mating success. In many lizard species, males are larger than females, whereas in others, females are the larger sex, suggesting that selection on fecundity has been stronger than sexual selection on males. As placental development or egg retention requires more space within the abdominal cavity, it has been suggested that females of viviparous lizards have larger abdomens or body size than their oviparous relatives. Thus, it would be expected that females of viviparous species attain larger sizes than their oviparous relatives, generating more biased patterns of SSD. We test these predictions using lizards of the genus Sceloporus. After controlling for phylogenetic effects, our results confirm a strong relationship between female body size and fecundity, suggesting that selection for higher fecundity has had a main role in the evolution of female body size. However, oviparous and viviparous females exhibit similar sizes and allometric relationships. Even though there is a strong effect of body size on female fecundity, once phylogenetic effects are considered, we find that the slope of male on female body size is significantly larger than one, providing evidence of greater evolutionary divergence of male body size. These results suggest that the relative impact of sexual selection acting on males has been stronger than fecundity selection acting on females within Sceloporus lizards.     

BODY SIZE AND SEXUAL SIZE DIMORPHISM IN MARINE IGUANAS FLUCTUATE AS A RESULT OF OPPOSING NATURAL AND SEXUAL SELECTION: AN ISLAND COMPARISON

Body size is often assumed to represent the outcome of conflicting selection pressures of natural and sexual selection. Marine iguana (Amblyrhynchus cristatus) populations in the Galápagos exhibit 10-fold differences in body mass between island populations. There is also strong sexual size dimorphism, with males being about twice as heavy as females. To understand the evolutionary processes shaping body size in marine iguanas, we analyzed the selection differentials on body size in two island populations (max. male mass 900 g in Genovesa, 3500 g in Santa Fé). Factors that usually confound any evolutionary analysis of body sizes—predation, interspecific food competition, reproductive role division—are ruled out for marine iguanas. We show that, above hatchlings, mortality rates increased with body size in both sexes to the same extent. This effect was independent of individual age. The largest animals (males) of each island were the first to die once environmental conditions deteriorated (e.g., during El Niños). This sex-biased mortality was the result of sexual size dimorphism, but at the same time caused sexual size dimorphism to fluctuate. Mortality differed between seasons (selection differentials as low as –1.4) and acted on different absolute body sizes between islands. Both males and females did not cease growth when an optimal body size for survival was reached, as demonstrated by the fact that individual adult body size phenotypically increased in each population under favorable environmental conditions beyond naturally selected limits. But why did marine iguanas grow “too large” for survival? Due to lek mating, sexual selection constantly favored large body size in males (selection differentials up to +0.77). Females only need to reach a body size sufficient to produce surviving offspring. Thereafter, large body size of females was less favored by fertility selection than large size in males. Resulting from these different selection pressures on male and female size, sexual size dimorphism was mechanistically caused by the fact that females matured at an earlier age and size than males, whereafter they constantly allocated resources into eggs, which slowed growth. The observed allometric increase in sexual size dimorphism is explained by the fact that the difference between these selective processes becomes larger as energy abundance in the environment increases. Because body size is generally highly heritable, these selective processes are expected to lead to genetic differences in body size between islands. We propose a common-garden experiment to determine the influence of genetic factors and phenotypic reaction norms of final body size.