Magnitude Of Sexual Size Dimorphism Research Articles

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.

Marine turtles are only minimally sexually size dimorphic, a pattern that is distinct from most nonmarine aquatic turtles.

Turtles have been prominent subjects of sexual size dimorphism (SSD) analyses due to their compact taxonomy, mating systems, and habitat diversity. In prior studies, marine turtles were grouped with fully aquatic non‐marine turtles (NMATs). This is interesting because it is well‐established that the marine environment imposes a distinct selective milieu on body form of vagile vertebrates, driven by convergent adaptations for energy‐efficient propulsion and drag reduction. We generated a comprehensive database of adult marine turtle body sizes (38,569 observations across all species), which we then used to evaluate the magnitude of SSD in marine turtles and how it compares to SSD in NMAT. We find that marine turtles are only minimally sexually size dimorphic, whereas NMAT typically exhibit female‐biased SSD. We argue that the reason for this difference is the sustained long‐distance swimming that characterizes marine turtle ecology, which entails significant energetic costs incurred by both sexes. Hence, the ability of either sex to allocate proportionately more to growth than the other is likely constrained, meaning that sexual differences in growth and resultant body size are not possible. Consequently, grouping marine turtles with NMAT dilutes the statistical signature of different kinds of selection on SSD and should be avoided in future studies.

Integrating growth and survival models for flexible estimation of size-dependent survival in a cryptic, endangered snake.

Estimates of demographic rates for animal populations and individuals have many applications for ecological and conservation research. In many animals, survival is size‐dependent, but estimating the form of the size–survival relationship presents challenges. For elusive species with low recapture rates, individuals’ size will be unknown at many points in time. Integrating growth and capture–mark–recapture models in a Bayesian framework empowers researchers to impute missing size data, with uncertainty, and include size as a covariate of survival, capture probability, and presence on‐site. If there is no theoretical expectation for the shape of the size–survival relationship, spline functions can allow for fitting flexible, data‐driven estimates. We use long‐term capture–mark–recapture data from the endangered San Francisco gartersnake (Thamnophis sirtalis tetrataenia) to fit an integrated growth–survival model. Growth models showed that females reach longer asymptotic lengths than males and that the magnitude of sexual size dimorphism differed among populations. The capture probability and availability of San Francisco gartersnakes for capture increased with snout–vent length. The survival rate of female snakes exhibits a nonlinear relationship with snout–vent length (SVL), with survival flat between 300 mm and 550 mm SVL before decreasing for females between 550 mm and 700 mm SVL. For male snakes, survival decreased for adult males >550 mm SVL. The survival rates of the smallest and largest San Francisco gartersnakes were highly uncertain because recapture rates were very low for these sizes. By integrating growth and survival models and using penalized splines, we found support for size‐dependent survival in San Francisco gartersnakes. Our results have applications for devising management activities for this endangered subspecies, and our methods could be applied broadly to the study of size‐dependent demography among animals.

Effects of Early Thermal Environment on Growth, Age at Maturity, and Sexual Size Dimorphism in Arctic Charr

The effects of early thermal environment on growth, age at maturity, and sexual size dimorphism in Arctic charr (Salvelinus alpinus) are investigated. This study is a 654-day long rearing trial split into two sequential experimental phases termed EP1 and EP2 and lasting 315 and 339 days, respectively. EP1 started at the end of the yolk sac stage when the experimental fish were divided into three groups and reared at different target temperatures (7, 10 and 12 °C). During EP2, all groups were reared at the same temperature (7–8 °C) until harvest (~1300 g). Growth rates increased with temperature from 7 to 12 °C, and at the end of EP1 the 12C group had 49.0% and 19.2% higher mean weight than groups 7C and 10C, respectively. Elevated early rearing temperatures were, however, found to cause precocious sexual maturation and reduce the long-term growth performance. At the end of EP2, the 7C group had 3.6% and 14.1% higher mean weight than 10C and 12C, respectively. Elevated early rearing temperatures had a much stronger effect on the maturity incidence of females, and while male-biased sexual size dimorphism (SSD) was found in all groups, the magnitude of SSD was positively associated with temperature.

Macroecological patterns of sexual size dimorphism in turtles of the world

Sexual size dimorphism (SSD) is a well-documented phenomenon in both plants and animals; however, the ecological and evolutionary mechanisms that drive and maintain SSD patterns across geographic space at regional and global scales are understudied, especially for reptiles. Our goal was to examine geographic variation of turtle SSD and to explore ecological and environmental correlates using phylogenetic comparative methods. We use published body size data on 135 species from nine turtle families to examine how geographic patterns and the evolution of SSD are influenced by habitat specialization, climate (annual mean temperature and annual precipitation) and climate variability, latitude, or a combination of these predictor variables. We found that geographic variation, magnitude and direction of turtle SSD are best explained by habitat association, annual temperature variance and annual precipitation. Use of semi-aquatic and terrestrial habitats was associated with male-biased SSD, whereas use of aquatic habitat was associated with female-biased SSD. Our results also suggest that greater temperature variability is associated with female-biased SSD. In contrast, wetter climates are associated with male-biased SSD compared with arid climates that are associated with female-biased SSD. We also show support for a global latitudinal trend in SSD, with females being larger than males towards the poles, especially in the families Emydidae and Geoemydidae. Estimates of phylogenetic signal for both SSD and habitat type indicate that closely related species occupy similar habitats and exhibit similar direction and magnitude of SSD. These global patterns of SSD may arise from sex-specific reproductive behaviour, fecundity and sex-specific responses to environmental factors that differ among habitats and vary systematically across latitude. Thus, this study adds to our current understanding that while SSD can vary dramatically across and within turtle species under phylogenetic constraints, it may be driven, maintained and exaggerated by habitat type, climate and geographic location.

Sex‐dependent responses of perch to changes in water clarity and temperature

AbstractRising temperatures and decreasing water transparency of lakes have strong wide ranging effects on fish. Fish responses to various changes in the environment are usually species‐dependent, but responses may also vary within species. In general, large individuals are considered to be more sensitive to environmental variation due to higher energy demand, than smaller individuals. Similarly, large individuals require more food to maintain bodily functions and are thus more sensitive to resource and food scarcity. These size‐specific responses to environmental gradients are also sex‐dependent in species that exhibit sexual size dimorphism (SSD). We studied in enclosures with short‐term experiments how rising temperatures and decreasing water transparency regulate the feeding rates of female and male European perch (Perca fluviatilis L.). To explore experimental results, we calculated perch SSD in nine lakes with varying environmental conditions using previously collected field data. The results of the experiments revealed that the combined effect of water transparency and temperature on the feeding rate of fish is gender‐dependent: feeding rate of females decreased more than that of males. The experimental results were also supported by field data that revealed a negative relation between water transparency and the magnitude of SSD in perch. Our results suggest that rising temperatures and decreasing water transparency may potentially decrease fish size in a sex‐dependent manner. As female size is one of the main demographic traits determining the reproductive success of a fish population, changing environments may have unexpected and far‐reaching consequences on fish population dynamics.

Correlated Evolution of Allometry and Sexual Dimorphism across Higher Taxa

Empirical evidence suggests that Rensch's rule of allometric scaling of male and female body size, which states that body size divergence is greater across males than across females of a clade, is not universal. In fact, quantitative genetic theory indicates that the sex under historically stronger directional selection will exhibit greater interspecific variance in size. Thus, the pattern of covariance between allometry of male and female body size and sexual size dimorphism (SSD) across related clades allows a test of this causal hypothesis for macroevolutionary trends in SSD. We compiled a data set of published body size estimates from the amphibians, a class with predominantly female-biased SSD, to examine variation in allometry and SSD among clades. Our results indicate that females become the more size-variant sex across species in a family as the magnitude of SSD in that family increases. This rejects Rensch's rule and implicates selection on females as a driver of both amphibian allometry and SSD. Further, when we combine our data into a single analysis of allometry for the class, we find a significant nonlinear allometric relationship between female body size and male body size. These data suggest that allometry changes significantly as a function of size. Our results illustrate that the relationship between female size and male size varies with both the degree of sexual dimorphism and the body size of a clade.

The Role of Sexual Selection and Conflict in Mediating Among-Population Variation in Mating Strategies and Sexually Dimorphic Traits in Sepsis punctum

The black scavenger fly Sepsis punctum exhibits striking among-population variation in the direction and magnitude of sexual size dimorphism, modification to the male forelimb and pre-copulatory behaviour. In some populations, male-biased sexual size dimorphism is observed; in other, less dimorphic, populations males court prior to mating. Such variation in reproductive traits is of interest to evolutionary biologists because it has the potential to limit gene flow among populations, contributing to speciation. Here, we investigate whether large male body size and modified forefemur are associated with higher male mating success within populations, whether these traits are associated with higher mating success among populations, and if these traits carry viability costs that could constrain their response to sexual selection. Flies from five distinct populations were reared at high or low food, generating high and low quality males. The expression of body size, forelimb morphology and courtship rate were each greater at high food, but high food males experienced higher mating success or reduced latency to first copulation in only one of the populations. Among populations, overall mating success increased with the degree of male-bias in overall body size and forelimb modification, suggesting that these traits have evolved as a means of increasing male mating rate. The increased mating success observed in large-male populations raises the question of why variation in magnitude of dimorphism persists among populations. One reason may be that costs of producing a large size constrain the evolution of ever-larger males. We found no evidence that juvenile mortality under food stress was greater for large-male populations, but development time was considerably longer and may represent an important constraint in an ephemeral and competitive growth environment.

Implications of Male and Female Contributions to Sexual Size Dimorphism for Inferring Behavior in the Hominin Fossil Record

Sexual dimorphism is commonly used to directly infer or support reconstructions of social behavior in early hominins. This is often done by comparing the magnitude of sexual size dimorphism to that seen in extant primates and extrapolating a likely social behavior. Such comparisons are of limited value, though, allowing only the inference of strong male–male competition when dimorphism is strong. Recent studies have begun to focus on the selective factors that impact female body size, and thereby size dimorphism. Considerations of changes in male and female size in the fossil record potentially allow insight into the meaning of changes in sexual dimorphism through time. To illustrate, I compare estimates of body mass dimorphism for four hominin taxa to assess changes in male and female size. Assuming that early Homo represents a single taxon, sexual size dimorphism increased in early Homo through an increase in male size, but was subsequently reduced through an increase in female size in Homo erectus. This would imply a significant increase in sexual selection acting on males in early Homo. An increase in female size with a loss of dimorphism in Homo erectus would imply a simultaneous shift in female optimal body size through selection for increased female fecundity, and/or an increase in female resource abundance, coupled with a shift in selection acting on male size. Although none of these inferences are certain, the exercise illustrates the potential for considering how dimorphism changes through time, rather than simply focusing on the magnitude of size dimorphism in isolation.

Rensch’s Rule in Large Herbivorous Mammals Derived from Metabolic Scaling

Rensch's rule, which states that the magnitude of sexual size dimorphism tends to increase with increasing body size, has evolved independently in three lineages of large herbivorous mammals: bovids (antelopes), cervids (deer), and macropodids (kangaroos). This pattern can be explained by a model that combines allometry, life-history theory, and energetics. The key features are that female group size increases with increasing body size and that males have evolved under sexual selection to grow large enough to control these groups of females. The model predicts relationships among body size and female group size, male and female age at first breeding, death and growth rates, and energy allocation of males to produce body mass and weapons. Model predictions are well supported by data for these megaherbivores. The model suggests hypotheses for why some other sexually dimorphic taxa, such as primates and pinnipeds (seals and sea lions), do or do not conform to Rensh's rule.

Sexual size dimorphism decreases with temperature in a blowfly, Chrysomya megacephala (Fabricius) (Diptera: Calliphoridae)

1. There is wide intra-specific variation in sexual size dimorphism (SSD). Much of this variation is probably as a result of sexual differences in the selective pressure on body size. However, environmental variables could affect males and females differently, causing variation in SSD. 2. We examined the effects of two temperatures (20 and 30 °C) on SSD in six populations of the blowfly, Chrysomya megacephala. 3. We found that body size increased with temperature in all the populations studied, and the sexes differed in phenotypic plasticity of body size in response to rearing temperature. This created substantial temperature-induced variation in SSD (i.e. sex × temperature interaction). Males were often smaller than females, but the degree of dimorphism was smaller at the higher temperature (30 °C) and larger at the lower temperature (20 °C). This change in SSD was not because of a gender difference in the effect of temperature on development time. Further studies should address whether this variation can be produced by adaptive canalisation of one sex against variation in temperature, or whether it may be a consequence of non-adaptive developmental differences between the sexes. 4. Although most studies assume that the magnitude of SSD is fixed within a species, the present study demonstrates that rearing temperature can generate considerable intra-specific variation in the degree of SSD.

Why are reproductively parasitic fish males so small?—influence of tactic-specific selection

Despite the wide prevalence of alternative reproductive tactics, little attention has been paid to why reproductively parasitic males are so small. In this study, we tackled this issue in a shell-brooding fish Lamprologus callipterus. Sneaky 'dwarf males' of this fish remain much smaller than bourgeois conspecifics throughout their life and employ a unique parasitic tactic, i.e. entering into a gastropod shell where a female is spawning, passing through the space between the female and shell wall and staying behind her to ejaculate throughout the spawning event. Here, we tested the prediction that they remain small to get past her through the shell spaces by interpopulation comparison. We showed, across populations, a negative allometry for sexual size dimorphism, an exponential increase of female size with an increase in shell size and a negative correlation between the magnitude of sexual size dimorphism and shell size. These results suggest that the inner spaces strongly regulate dwarf male size. We conclude that the small bodies of dwarf males arise from adaptation to their unique reproductive behaviour.

SEX-SPECIFIC SELECTION AND INTRASPECIFIC VARIATION IN SEXUAL SIZE DIMORPHISM

Sexual size dimorphism (SSD) is thought to evolve due to sex differences in selection on body size, but it is largely unknown whether intraspecific variation in SSD reflects differences in sex-specific selection among populations. We addressed this question by comparing viability selection between two island populations of the brown anole lizard (Anolis sagrei) that differ in the magnitude of male-biased SSD. On both islands, females experienced stabilizing selection favoring intermediate size whereas males experienced directional selection favoring larger size. Thus, sex-specific selection matched the overall pattern of male-biased SSD, but population differences in the magnitude of SSD were not associated with local differences in selection. Rather, population differences in SSD appear to result from underlying differences in the environmental potential for a rapid growth, coupled with sex-specific phenotypic plasticity. Males grew more slowly on the island with low SSD whereas growth of females did not differ between islands. Both sexes had substantially lower mass per unit length on the island with low SSD, suggesting that they were in a relatively poorer energetic condition. We propose that this energetic constraint disproportionately impacts growth of males due to their greater absolute energy requirements, thus driving intraspecific variation in SSD.

A Phylogenetic Analysis of Sexual Size Dimorphism in Turtles

Sexual size dimorphism (SSD) is widespread in a variety of vertebrate taxa, and its presence has been associated with a range of evolutionary and ecological factors. Turtles, as a lineage, have been shown to include species that exhibit both forms of SSD—male-biased (male larger) and female-biased (female larger)—as well as species that do not exhibit SSD (male size ≅ female size). Previous work has suggested that factors affecting the direction and magnitude of SSD in turtles may include habitat preference and fecundity; these correlations, however, have not been tested in a phylogenetic context. To provide a framework for evaluating potential correlates with the magnitude and direction of SSD in turtles, we examined the phylogenetic distribution of SSD in this clade. Published sources were surveyed for data on the direction and magnitude of SSD, carapace length, clutch size, egg size, and preferred habitat (terrestrial vs. nonterrestrial). Phylogenies were compiled from multiple published sources to develop a phylogeny for all taxa (n = 82) for which SSD data were assembled. The phylogenetic distribution of SSD was then evaluated using parsimony-based methods. These analyses indicated that the ancestral condition for extant turtles was female-biased SSD. Phylogenetic analyses conducted on subsets of the data (concentrated-changes tests, independent contrasts) indicated that changes in SSD state were not correlated with changes in habitat preference. Both male and female carapace length were found to be correlated with traits related to fecundity (egg size, clutch size), but the degree of dimorphism was not found to be correlated with these traits. A significant negative correlation between egg size and clutch size was also evident in turtles, as predicted by life-history theory.

Variation in the magnitude of sexual size dimorphism in nestling Coal Tits (Periparus ater)

The magnitude of sexual size dimorphism can be affected by sex differences in environmental sensitivity early in ontogeny that result in differential growth rates of male and female nestlings. Here, the larger sex might either be more sensitive because of higher food demands or less sensitive due to greater competitive ability. When environmental conditions deteriorate during the breeding season, this “environmental stress” hypothesis predicts differential seasonal declines in the performance of male and female offspring. Based on a sample of molecularly sexed Coal Tit (Periparus ater) nestlings from 2 years, we investigated sexual size dimorphism in body mass, condition (i.e. size-corrected mass), tarsus and wing length and whether its magnitude changed from early to late broods. Male offspring were heavier, larger (in terms of tarsus and wing length) and had higher size-corrected mass than their female nest mates (the same was evident in adult breeders). In 2002 (the year with the longer effective breeding season), body mass and condition declined with progressing hatching date and this effect was significantly more pronounced in male than in female nestlings. There was also a seasonal decline in male wing length, while female wing length remained relatively constant, which resulted in males having shorter wings than females in late broods. Tarsus length was unaffected by time of breeding, except that the difference between males and females was relatively smaller in late (i.e. second) broods in 2002. While these results are in accordance with the idea of an increased environmental sensitivity of the larger males, confounding effects of sex-differential hatching order cannot be ruled out.

Sexual size dimorphism in a Drosophila clade, the D. obscura group

The Drosophila obscura clade consists of about 41 species, of which 20 were used for analyses of wing and thorax length. Our primary goal was to investigate the magnitude of sexual size dimorphism (SSD) of these traits within this clade and to test Rensch's Rule [when females are larger than males, SSD (e.g., female/male ratio) should decrease with body size]. Our secondary goal was methodological and involved evaluating for these flies alternative measures of SSD (female/male ratio, female/male absolute difference, female/male relative difference), developing a bootstrap method to estimate the magnitude of intraspecific variation in SSD, and applying a new method of estimating allometric relationships that is phylogenetically based and incorporates error variance in both traits. All indices of SSD were strongly correlated for both size traits. Nevertheless, female/male ratio is the best index here: it is easily interpretable and essentially independent of size. For both traits, SSD (F/M) varied interspecifically, showed a strong phylogenetic signal, but did not differ for the main phylogenetic subgroups or correlate with latitude. Factors underlying variation in SSD in this clade are elusive and might include genetic drift. SSD (wing) tended to decrease with increasing size, as predicted by Rensch's Rule, though not consistently so. SSD (thorax) was unrelated to size. However, analysis of published data for thorax length of Drosophila spp. ( N=42) with a larger size range showed that SSD decreased significantly with increasing size (consistent with Rensch's Rule), suggesting our ability to detect SSD-size relations in the D. obscura data may be limited by low statistical power.

Sexual selection explains Rensch's rule of size dimorphism in shorebirds.

Sexual size dimorphism shows a remarkably widespread relationship to body size in the animal kingdom: within lineages, it decreases with size when females are the larger sex, but it increases with size when males are the larger sex. Here we demonstrate that this pattern, termed Rensch's rule, exists in shorebirds and allies (Charadriides), and it is determined by two components of sexual selection: the intensity of sexual selection acting on males and the agility of the males' display. These effects are interactive so that the effect of sexual selection on size dimorphism depends on male agility. As a control, we also examine dimorphism in bill length, which is a functionally selected trait. As such, dimorphism in bill length neither exhibits Rensch's rule nor is associated with sexual selection and display. Our results show that variation among taxa in the direction and magnitude of sexual size dimorphism, as manifested as Rensch's rule, can be explained by the interaction between the form and strength of sexual selection acting on each sex in relation to body size.

Reinventing the Wheel: Analysis of Sexual Dimorphism in Body Size

Sexual dimorphism in body size has received increasing scientific interest during the past decade. In most of the papers on the subject, the magnitude of sexual size dimorphism is expressed as some conversion of body size ratio. There are several possible flaws in using such ratios, however, the most serious one concers the scaling of body size and size differences between sexes. Using ratios may easily lead to wrong conclusions. In this paper we emphasize some fallacies related to ratios and present a less biased method, based on residuals of linear regression. This method is suggested to be used in comparative analyses of sexual size dimorphism.

THE EVOLUTION OF FEMALE BODY SIZE IN RED-WINGED BLACKBIRDS: THE EFFECTS OF TIMING OF BREEDING, SOCIAL COMPETITION, AND REPRODUCTIVE ENERGETICS.

We examined opposing selective forces on female body size in the sexually dimorphic red-winged blackbird: social competition favoring larger females, and energetic advantages favoring smaller females. Downhower proposed that selection might drive female birds to be smaller than the optimum for survival, if smaller females were able to exceed their energetic requirements for self-maintenance earlier in the season and therefore breed earlier. Since in most birds the earliest breeders fledge the most young, this could favor the evolution of smaller female size, and therefore contribute to the magnitude of sexual size dimorphism in these birds. We tested this hypothesis in 1987 and 1988 by comparing the size and breeding date of female red-winged blackbirds. Consistent with our preditions, early-nesting females had much higher nesting success, but contrary to prediction, larger females bred earlier. We then examined the effects of female size on competition. If large females have an advantage in social competition, and if competition influences breeding date and reproductive success, then larger females might breed earlier. Primary females, the first females to arrive and nest on a territory, were more aggressive than lower ranked females; more aggressive females settled on better territories and laid earlier than less aggressive females; and larger females were more aggressive. Social competition between females may therefore favor large females. Finally, we tested the prediction that selection favoring large females might be limited by energetic constraints on large females. We found that large females had less fat than small females during breeding, and that the levels of fat that females of a given size carried affected breeding date and egg size. Therefore, social competition may favor large females, but reproductive energetics favoring smaller females may constrain selection for large female body size.

Sexual Size Dimorphism of Bird-Eating Hawks: The Effect of Prey Vulnerability

Among predatory birds, magnitude of sexual size dimorphism increases progressively from carrion-eating and invertebrate-eating species through fish-eating and mammal-eating species to a high in bird-eating species. Analysis of raptor hunting success demonstrates that the relation between hunting success and prey type is exactly opposite the relation between prey type and degree of dimorphism: raptors that prey upon birds experience more difficulty in the successful capture of their prey than raptors that prey upon fish, reptiles, or invertebrates. Consequently, a new hypothesis based upon prey vulnerability is introduced to explain differences in degree of raptor dimorphism. The prey vulnerability hypothesis maintains that lower encounter rates and capture successes resulting from the highly developed predator detection and escape abilities of avian prey may lower the availability of avian prey in comparison to nonavian prey. This would result in bird-eating raptors receiving a lower net energy intake per unit time spent foraging than non-bird-eating raptors. The prey vulnerability hypothesis supplements intersexual food competition or sexual role division hypotheses by presenting an alternative mechanism based upon prey availabilities (not prey abundances) for how differences among prey types may have led to differences in degree of raptor size dimorphism.