Sexual size dimorphism and male combat in snakes

Summary This paper reviews published literature on snakes to test the hypothesis that large male size, relative to female size, evolves because of the advantage it confers in male combat. Analysis of the data reveals a high correlation between the occurrence of male combat, and sexual dimorphism in which the male is the larger sex. This correlation holds (i) within the total sample of snake species (n=224), (ii) within the family Colubridae (n=134), and (iii) in a comparison between the eight families of snakes for which data are available. These results strongly support the hypothesis that large male size is an adaptation to intrasexual competition. The analysis also shows that females are larger than males in about 66% of snake species, that male combat is known in only about 15% of species, and that both sexual size dimorphism and the incidence of male combat tend to be distributed along taxonomic lines.     

Sexual size dimorphism and sexual selection in turtles (order testudines)

Summary This paper combines published and original data on sexual size dimorphism, reproductive behavior, and habitat types in turtles. Our major finding is that observed patterns of sexual size dimorphism correlate with habitat type and male mating strategy. (1) In most terrestrial species, males engage in combat with each other. Males typically grow larger than females. (2) In semiaquatic and bottom-walking aquatic species, male combat is less common, but males often forcibly inseminate females. As in terrestrial species, males are usually larger than females. (3) In truly aquatic species, male combat and forcible insemination are rare. Instead, males utilize elaborate precoital displays, and female choice is highly important. Males are usually smaller than females.We interpret these correlations between sexual behavior and size dimorphism in terms of sexual selection theory: males are larger than females when large male size evolves as an adaptation to increase success in male combat, or to enable forcible insemination of females. In contrast, males are usually smaller than females where small size in males evolves to increase mobility (and hence, ability to locate females), or because selection for increased fecundity may result in increased female size. In turtle species with male combat or forcible insemination, the degree of male size superiority increases with mean species body size.     

Mating success in the spittlebug Cercopis sanguinolenta (Scopoli, 1763) (Homoptera, Cercopidae): the role of body size and mobility

In insects, a sexual size dimorphism commonly occurs, with larger females. However, as a deviation from this general rule, larger males are found in some species. In these species often sexual selection for large males has been presumed. The spittlebug Cercopis sanguinolenta exhibits a distinct sexual size dimorphism with larger males. Mating behaviour was studied in a field population in respect to mating success of males and females. The aim of this study was to examine the mechanisms that lead to the observed non-random mating pattern. The results showed a mating pattern without size-assortative mating. A correlation was found between mating success and body size in males. In females no such correlation was found. The mobility of males depends on their body size and mobility is high only when females are present. However, in an analysis of covariance it was found that male mating success is not correlated with mobility, when controlled for body size. The mating system of the spittlebug was classified as scramble competition polygyny. Electronic Publication     

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.     

拟环纹豹蛛雄蛛触肢上黑白相间毛饰物的功能

狼蛛科雄蛛附肢上多样化的饰装往往与求偶行为相偶联,这些特殊的饰装通常被认为是雌性选择的结果。拟环纹豹蛛Pardosa pseudoannulata,属狼蛛科豹蛛属,雄蛛触肢胫节密被白毛,跗舟密被黑毛,具有典型的性二型现象;同时,只有成熟的雄蛛才展现触肢黑白相间的毛饰物。推测拟环纹豹蛛雄蛛触肢这种黑白相间的毛饰物可能在物种识别中具重要作用。在室内我们拟通过涂抹操作对拟环纹豹蛛雄蛛触肢黑白相间毛饰物的功能进行分析。实验分为4组,分别是对照组(A组,雄蛛不做任何处理)、雄蛛触肢白色胫节全部涂成黑色(B组)、雄蛛触肢黑色跗舟全部涂成白色(C组)和雄蛛触肢的黑色跗舟被涂成黑色(D组),然后采用雌雄配对进行求偶交配行为测定。实验结果表明,B组雄蛛的交配成功率显著低于A、C和D组的雄蛛,而后3组雄蛛的交配成功率无显著差异。相反,B组雄蛛被雌蛛相食百分率显著高于其它3组。可见拟环纹豹蛛雄蛛触肢上黑白相间毛饰物,尤其是其胫节上的白色饰物在雌蛛种间识别中起重要作用。     

Dishonest signals of strength in male slender crayfish (Cherax dispar) during agonistic encounters

Many animals resolve disputes without combat by displaying signals of potential strength during threatening displays. Presumably, competitors use each other's displays to assess their relative strengths, and current theory predicts that these signals of strength should generally be honest. We tested this prediction by investigating the relationships among morphology, performance, and social dominance in males of the slender crayfish Cherax dispar. Crayfish routinely use their enlarged front claws (chelae) for both intimidation and fighting, making this species ideal for studying the honesty of weapon size. We evaluated five competing models relating morphological and physiological traits to dominance during paired competitive bouts. Based on the best model, larger chelae clearly resulted in greater dominance; however, chela strength had no bearing on dominance. Thus, displays of chela size were dishonest signals of strength, and the enlarged chelae of males seemingly function more for intimidation than for fighting. In addition, an analysis of the performance of isolated chela muscle showed that muscle from male crayfish produced only half the force that muscle from female crayfish produced (236.6+/-26.4 vs. 459.5+/-71.6 kN m(-2)), suggesting that males invest more in developing larger chelae than they do in producing high-quality chela muscle. From our studies of crayfish, we believe dishonest signaling could play a greater role in territorial disputes than previously imagined.     

Male mating success,sexual size dimorphism,and site fidelity in two species of Malacoctenus (Labrisomidae)

Synopsis In both Malacoctenus hubbsi and Malacoctenus macropus, males defended preferred oviposition sites from both other males and potential egg predators. In M. hubbsi, adult females were larger than adult males. Larger M. hubbsi males were not associated with territory parameters that were correlated with higher mating success, and male size was not correlated with mating success. Male size did affect mating success when territory parameters were statistically controlled for, but the failure of large males to associate with better territories eliminated any mating advantage for larger males. In M. macropus, males are larger than females. Larger males defended preferred oviposition sites, and had higher mating success than did smaller males. Male M. macropus also had much higher site fidelity than male M. hubbsi. These results suggest that the evolution of the differences in site fidelity and sexual size dimorphism between these two species may be due to sexual selection acting differentially in these two species.     

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.     

Seasonal change in sexual size dimorphism of the major cheliped in the hermit crab <Emphasis Type="Italic">Pagurus minutus</Emphasis>

Sexual size dimorphism is a common phenomenon in the animal kingdom, and its seasonal change has been reported in some species that possess traits dimorphic only in males and specialized for male mating success. However, few studies have examined seasonal change in sexual dimorphism of traits possessed by both sexes. Here, we examined the reproductive biology of the hermit crab Pagurus minutus, at a sandflat in the Waka River estuary, Japan, with special reference to seasonal changes in sexual dimorphism of the large claw (major cheliped) size by conducting population and precopulatory guarding-pair sampling. Previous investigation demonstrated that the major cheliped is used as a weapon, and its size, more than body size, determines the winner in male–male contests of this species. We found ovigerous females from November to April, peaking in January, when 80% of females were ovigerous. Sexual size dimorphism of the major cheliped was observed; the degree of dimorphism increased in the reproductive season, when only males possessed an enlarged major cheliped. In addition, in the reproductive season, precopulatory guarding males had a larger body and larger relative size of the major cheliped than did solitary males, although the major cheliped size in guarding males seemed to reach an upper limit. These results suggest that seasonal change in sexual dimorphism of the major cheliped size in P. minutus strongly reflects sexual selection favoring the development of this natural weaponry, and that the degree of the dimorphism might be limited through natural selection.     

Sexual dimorphism in the tusked frog, Adelotus brevis (Anura:Myobatrachidae): the roles of natural and sexual selection

At least two adaptive processes can lead to the evolution of sexual dimorphism: sexual selection (e.g. male-male combat) or natural selection (e.g. dietary divergence). We investigated the adaptive significance of a distinctive pattern of sexual dimorphism in a south-eastern Australian frog, Adelotus brevis. Male Adelotus grow larger than female conspecifics, have larger heads relative to body size, and have large paired projections (‘tusks’) in the lower jaw. All of these traits are rare among anurans. We quantified the degree of dimorphism in Adelotus, and gathered data on diets and mating systems of this species to evaluate the possible roles of sexual selection and dietary divergence in favoring die evolution of these sexually dimorphic traits. Analysis of prey items in alimentary tracts revealed significant sex differences in prey types. For example, females ate proportionally more arthropods and fewer molluscs than did males. However, this difference is likely to be a secondary consequence of habitat differences between the sexes (due in turn to their different reproductive roles) rather than a selective force for the evolution of sexual dimorphism. Calling males spend their time in moist habitats where pondsnails are abundant, whereas females are more often encountered in the drier arthropod-rich woodlands. A three-year behavioural ecology study on a field population revealed that reproductive males engage in agonistic interactions, with the sexually dimorphic tusks used to attack rivals. Larger body size contributed to male reproductive success. Small males were excluded from calling sites and, among the calling males, larger animals had higher reproductive success (numbers of matings) than did smaller individuals. Hence, the atypical pattern of sexual dimorphism in Adelotus brevis seems to have resulted from sexual selection for larger body size and tusk size in males, in the context of male-male agonistic behaviour, rather than natural selection for ecological divergence between the sexes.     

Territoriality and male-biased sexual size dimorphism in Argia reclusa (Odonata: Zygoptera)

In Odonata, many species present sexual size dimorphism (SSD), which can be associated with male territoriality in Zygoptera. We hypothesized that in the territorial damselfly Argia reclusa, male–male competition can favor large males, and consequently, drive selection pressures to generate male-biased SSD. The study was performed at a small stream in southeastern Brazil. Males were marked, and we measured body size and assessed the quality of territories. We tested if larger territorial males (a) defended the best territories (those with more male intrusions and visiting females), (b) won more fights, and (c) mated more. Couples were collected and measured to show the occurrence of sexual size dimorphism. Results indicated that males are larger than females, and that territorial males were larger than non-territorial males. Larger territorial males won more fights and defended the best territories. There was no difference between the mating success of large territorial and small non-territorial males. Although our findings suggest that male territoriality may play a significant role on the evolution of sexual size dimorphism in A. reclusa, we suggest that other factors should also be considered to explain the evolution of SSD in damselflies, since non-territorial males are also capable of acquiring mates.     

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.     

Ecological context and the importance of body and gnathopod size for pairing success in two amphipod ecomorphs

Ecological context generates interspecific variation in mating behavior by imposing differential constraints on the action of sexual selection, but operation of these constraints in nature is not well understood. We used field and laboratory studies to examine the importance of body size and size of sexually dimorphic appendages, the gnathopods, for pairing success in two freshwater amphipod species within the Hyalella azteca species complex. We focused on a large-bodied species found in habitats where ecological factors tend to favor large body size, and a small-bodied species in habitats where small body size is favored by size-selective predation. A field study indicated that although male pairing success was greater for larger males in both species, pairing success increased throughout the range of variation in male size in the large species, whereas, in the small species, pairing success was low for smaller individuals, but roughly constant across intermediate to larger sizes. A laboratory mate choice experiment was consistent with the field study, finding that females of the large species exhibited a preference for larger males that was independent of absolute male size, but females of the small species were indifferent when choosing between males of intermediate to larger size. Species also differed in the direction of sexual size dimorphism in the field, with males being the larger sex in the large species but the smaller sex in the small species, a pattern consistent with the species differences in mate preference. Large gnathopod size relative to body size was associated with enhanced pairing success across all body sizes for the large species, but, in the small species, large gnathopod size enhanced pairing success only in smaller males. Species differences in mating behavior appear consistent with change driven by differing forms of the interaction between sexual and natural selection.     

The evolution of performance-based male fighting ability in Caribbean Anolis lizards

Despite the empirical and theoretical attention paid to the role of sexual signals in resolving agonistic interactions between conspecific males, few studies have applied a comparative perspective, particularly across species that vary in combat intensity. We investigated the relative roles of a male sexual signal (dewlap size) and whole-organism performance capacity (bite force) on male combat outcomes in nine species of Caribbean Anolis lizards that differ markedly in territoriality, as indicated by sexual size dimorphism. We found that (1) dewlap size was generally an honest signal of bite force in dimorphic but not less dimorphic species; (2) maximum bite force consistently predicted male combat success in dimorphic but not less dimorphic species; (3) in contrast to a priori predictions, dewlap size significantly predicted male combat success in less dimorphic but not dimorphic species; and (4) the incidence of biting but not dewlapping increases as species become more dimorphic. These findings suggest that more dimorphic (and hence more territorial) species escalate to biting during fights more readily compared with less territorial species. The ecological and behavioral qualities of species may therefore modify both the shape and the size of sexually selected traits as well as the nature of the information those traits convey.     

The Role of Body Size in Mating Interactions of the Sexually Cannibalistic Fishing Spider Dolomedes triton

Some arachnids display extreme sexual size dimorphism (SSD) with adult females being several times larger than adult males. One explanation for SSD in species that exhibit pre‐copulatory sexual cannibalism (female attack, kill and consumption of the male prior to mating) is that smaller males may be less likely victims of predatory attacks by females. However, in some sexually cannibalistic species SSD is relatively moderate (i.e. males are similar in size to females) suggesting benefits of large male body size. Here, I report the results of an experiment designed to explore the ramifications of body size in mating interactions of the sexually cannibalistic, North American fishing spider (Dolomedes triton). Results suggest that male size does not influence courtship behavior, the likelihood of being attacked, or the male's ability to secure a mounting. However, large males were superior at gaining copulations once mounted. Sexual cannibalism may also be predicated on female size. Female condition (mass/cephalothorax area) did not explain any of these behaviors from the copulatory sequence, however, females with a smaller cephalothorax area were more likely to attack courting males. Finally, analysis of the ratio of female size to male size showed that when SSD is weak males are more likely to escape attacks and mate successfully. Results are discussed in light of several hypotheses for sexual cannibalism, and the benefits of large male body size illustrated here are put forth as potential explanations for the relatively moderate extent of SSD found in this sexually cannibalistic species.     

Sexual selection on skeletal shape in Carnivora

Lifetime reproductive success of males is often dependent upon the ability to physically compete for mates. However, species variation in social structure leads to differences in the relative importance of intraspecific aggression. Here, we present a large comparative dataset on sexual dimorphism in skeletal shape in Carnivora to test the hypotheses that carnivorans exhibit sexual dimorphism in skeletal anatomy that is reflective of greater specialization for physical aggression in males relative to females and that this dimorphism is associated with the intensity of sexual selection. We tested these hypotheses using a set of functional indices predicted to improve aggressive performance. Our results indicate that skeletal shape dimorphism is widespread within our sample. Functional traits thought to enhance aggressive performance are more pronounced in males. Phylogenetic model selection suggests that the evolution of this dimorphism is driven by sexual selection, with the best‐fitting model indicating greater dimorphism in polygynous versus nonpolygynous species. Skeletal shape dimorphism is correlated with body size dimorphism, a common indicator of the intensity of male–male competition, but not with mean body size. These results represent the first evidence of sexual dimorphism in the primary locomotor system of a large sample of mammals.     

`Little Chipmunk' Syndrome? Male Body Size and Dominance in Captive Yellow‐Pine Chipmunks (Tamias amoenus)

In mammals, large males are often assumed to have higher mating success because they have greater success at contest competition. This relationship is often used to explain the prevalence of male-biased sexual size dimorphism in mammals. However, in many small vertebrates, large individuals are not always dominant. Using staged dyadic encounters, we examined the relationship between male body size and social dominance in captive male yellow-pine chipmunks ( Tamias amoenus ), a species with female-biased sexual size dimorphism. The yellow-pine chipmunk has a mating system in which males participate in mating chases and dominant males may have an advantage in acquiring matings with oestrous females. Captive male chipmunks were aggressive in only 28% of 144 paired encounters; however, several lines of evidence indicated that smaller chipmunks were dominant over large chipmunks: (1) small males were dominant in more dyads than large males; (2) within dyads, dominant males were smaller than subordinate males; and (3) small males performed more aggressive behaviour than large males. These results are not consistent with the prediction that large males are typically dominant. If large chipmunks are able to gain matings with females because of qualities other than dominance (such as the ability to successfully find and/or chase receptive females), then the costs of aggression to large chipmunks may outweigh any potential benefits. Small males, but not large males, may improve their mating success by being aggressive.     

Sexual selection for male mobility in a giant insect with female-biased size dimorphism

Female-biased size dimorphism, in which females are larger than males, is prevalent in many animals. Several hypotheses have been developed to explain this pattern of dimorphism. One of these hypotheses, the mobility hypothesis, suggests that female-biased size dimorphism arises because smaller males are favored in scramble competition for mates. Using radiotelemetry, we assessed the mobility hypothesis in the Cook Strait giant weta (Deinacrida rugosa), a species with strong female-biased size dimorphism, and tested the prediction that male traits promoting mobility (i.e., longer legs, smaller bodies) are useful in scramble competition for mates and thus promote reproductive success. Our predictions were supported: males with longer legs and smaller bodies exhibited greater mobility (daily linear displacement when not mating), and more mobile males had greater insemination success. No phenotypic traits predicted female mobility or insemination success. In species with female-biased size dimorphism, sexual selection on males is often considered to be weak compared to species in which males are large or possess weaponry. We found that male giant weta experience sexual selection intensities on par with males of a closely related harem-defending polygynous species, likely because of strong scramble competition with other males.     

Is diversification in male reproductive traits driven by evolutionary trade-offs between weapons and nuptial gifts?

Many male animals have evolved exaggerated traits that they use in combat with rival males to gain access to females and secure their reproductive success. But some male animals invest in nuptial gifts that gains them access to females. Both these reproductive strategies are costly in that resources are needed to produce the weapon or nuptial gift. In closely related species where both weapons and nuptial gifts are present, little is known about the potential evolutionary trade-off faced by males that have these traits. In this study, we use dobsonflies (order Megaloptera, family Corydalidae, subfamily Corydalinae) to examine the presence and absence of enlarged male weapons versus nuptial gifts within and among species. Many dobsonfly species are sexually dimorphic, and males possess extremely enlarged mandibles that they use in battles, whereas in other species, males produce large nuptial gifts that increase female fecundity. In our study, we show that male accessory gland size strongly correlates with nuptial gift size and that when male weapons are large, nuptial gifts are small and vice versa. We mapped weapons and nuptial gifts onto a phylogeny we constructed of 57 species of dobsonflies. Our among-species comparison shows that large nuptial gift production evolved in many species of dobsonfly but is absent from those with exaggerated weapons. This pattern supports the potential explanation that the trade-off in resource allocation between weapons and nuptial gifts is important in driving the diversity of male mating strategies seen in the dobsonflies, whereas reduced male–male competition in the species producing large spermatophores could be an alternative explanation on their loss of male weapons. Our results shed new light on the evolutionary interplay of multiple sexually selected traits in animals.