Variation In Sexual Size Dimorphism Research Articles

Spotted Hyena skull size variation across geography favors the energetic equivalence rule over Bergmann’s Rule

Abstract Much historic work has focused on establishing geographical and ecological rules that broadly explain patterns in size variation. We examined geographic variation in Spotted Hyena skull size using geometric morphometrics and spatial statistics. We quantified size variation and sexual size dimorphism of the skull, and evaluated the influence of temperature, precipitation, land cover type, and population density on skull size. We found that female spotted hyenas are slightly larger on average than males. Our analysis of regional differences did not indicate geographic variation in sexual size dimorphism. Skull size of Spotted Hyenas varies with geography but does not adhere to Bergmann’s Rule. The smallest individuals of both sexes occur between −5.00° and 10.00° latitude and east of 28.50° longitude, with larger individuals being found elsewhere. Although Spotted Hyena skull size co-varies in some views with such variables as habitat type and climate indicators, skull size in this species most strongly co-varies with population density. The highest population densities are associated with the smallest skull size, possibly reflecting a relationship between high population density and access to resources. These results suggest that geographic variation in Spotted Hyena skull size is better explained by the energetic equivalence rule than Bergmann’s Rule.

The macroevolution of sexual size dimorphism in birds

Abstract There is considerable variation of sexual size dimorphism (SSD) in body mass among animal groups, yet the drivers of interspecific variation in SSD are still poorly understood. Possible mechanisms have been suggested, including sexual selection, selection for fecundity in females, niche divergence between sexes, and allometry, yet their relative importance is still poorly understood. Here, we tested predictions of these four hypotheses in different avian groups using a large-scale dataset on SSD of body mass for 4761 species. Specifically, we estimated the probability of transition between male- and female-biased SSD, tested for differences in evolutionary rates of body mass evolution for males and females, and assessed the potential ecological and spatial correlates of SSD. Our results were consistent with the sexual selection, fecundity, and niche divergence hypotheses, but their support varied considerably among avian orders. In addition, we found little evidence that the direction of SSD affected the evolution of male or female body mass, and no relationship was detected between SSD and environmental predictors (i.e. temperature and precipitation seasonality, productivity, species richness, and absolute latitude). These results suggest that avian evolution of SSD is likely to be multifactorial, with sexual selection, fecundity, and niche divergence playing important roles in different avian orders.

Revisiting niche divergence hypothesis in sexually dimorphic birds: Is diet overlap correlated with sexual size dimorphism?

The evolution of sexual size dimorphism (SSD) is a long-standing topic in evolutionary biology, but there is little agreement on the extent to which SSD is driven by the different selective forces. While sexual selection and fecundity selection have traditionally been proposed as the two leading hypotheses, SSD may also result from natural selection through mechanisms such as sexual niche divergence, which might have reduced resource competition between sexes. Here, we revisited the niche divergence hypothesis by testing the relationship between the sexual overlap in diet and SSD of 56 bird species using phylogenetic comparative analyses. We then assessed how SSD variation relates to the three main hypotheses: sexual selection, fecundity selection, and sexual niche divergence using phylogenetic generalized least squares (PGLS). Then, we compared sexual selection, fecundity selection and niche divergence selection as SSD drivers through phylogenetic confirmatory path analyses to disentangle the possible causal evolutionary relationships between SSD and the three hypotheses. Phylogenetic generalized least squares showed that SSD was negatively correlated with diet overlap, that is, the greater the difference in body size between males and females, the less diet overlap. As predicted by sexual selection theory, the difference in body size between sexes was higher in polygynous species. Confirmatory phylogenetic path analyses suggested that the most likely evolutionary path might include the mating system as a main driver in SSD and niche divergence as a result of SSD. We found no evidence of a role of fecundity selection in the evolution of female-biased SSD. Our study provides evidence that sexual selection has likely been the main cause of SSD and that dietary divergence is likely an indirect effect of SSD.

The origins and drivers of sexual size dimorphism in sharks.

While sexual size dimorphism (SSD) is abundant in nature, there is huge variation in both the intensity and direction of SSD. SSD results from a combination of sexual selection for large male size, fecundity selection for large female size and ecological selection for either. In most vertebrates, it is variation in the intensity of male-male competition that primarily underlies variation in SSD. In this study, we test four hypotheses regarding the adaptive value of SSD in sharks-considering the potential for each of fecundity, sexual, ecological selection and reproductive mode as the primary driver of variation in SSD between species. We also estimate past macroevolutionary shifts in SSD direction/intensity through shark phylogeny. We were unable to find evidence of significant SSD in early sharks and hypothesise that SSD is a derived state in this clade, that has evolved independently of SSD observed in other vertebrates. Moreover, there is no significant relationship between SSD and fecundity, testes mass or oceanic depth in sharks. However, there is evidence to support previous speculation that reproductive mode is an important determinant of interspecific variation in SSD in sharks. This is significant as in most vertebrates sexual selection is thought to be the primary driver of SSD trends, with evidence for the role of fecundity selection in other clades being inconsistent at best. While the phylogenetic distribution of SSD among sharks is superficially similar to that observed in other vertebrate clades, the relative importance of selective pressures underlying its evolution appears to differ.

Y-Linked Copy Number Polymorphism of Target of Rapamycin Is Associated with Sexual Size Dimorphism in Seed Beetles.

The Y chromosome is theorized to facilitate evolution of sexual dimorphism by accumulating sexually antagonistic loci, but empirical support is scarce. Due to the lack of recombination, Y chromosomes are prone to degenerative processes, which poses a constraint on their adaptive potential. Yet, in the seed beetle, Callosobruchus maculatus segregating Y linked variation affects male body size and thereby sexual size dimorphism (SSD). Here, we assemble C. maculatus sex chromosome sequences and identify molecular differences associated with Y-linked SSD variation. The assembled Y chromosome is largely euchromatic and contains over 400 genes, many of which are ampliconic with a mixed autosomal and X chromosome ancestry. Functional annotation suggests that the Y chromosome plays important roles in males beyond primary reproductive functions. Crucially, we find that, besides an autosomal copy of the gene target of rapamycin (TOR), males carry an additional TOR copy on the Y chromosome. TOR is a conserved regulator of growth across taxa, and our results suggest that a Y-linked TOR provides a male specific opportunity to alter body size. A comparison of Y haplotypes associated with male size difference uncovers a copy number variation for TOR, where the haplotype associated with decreased male size, and thereby increased sexual dimorphism, has two additional TOR copies. This suggests that sexual conflict over growth has been mitigated by autosome to Y translocation of TOR followed by gene duplications. Our results reveal that despite of suppressed recombination, the Y chromosome can harbor adaptive potential as a male-limited supergene.

Genetic Variation in Sexual Size Dimorphism Is Associated with Variation in Sex-Specific Plasticity in Drosophila.

AbstractThe difference in body size between females and males, or sexual size dimorphism (SSD), is ubiquitous, yet we have a poor understanding of the developmental genetic mechanisms that generate it and how these mechanisms may vary within and among species. Such an understanding of the genetic architecture of SSD is important if we are to evaluate alternative models of SSD evolution, but the genetic architecture is difficult to describe because SSD is a characteristic of populations, not individuals. Here, we overcome this challenge by using isogenic lineages of Drosophila to measure SSD for 196 genotypes. We demonstrate extensive genetic variation for SSD, primarily driven by higher levels of genetic variation for body size among females than among males. While we observe a general increase in SSD with sex-averaged body size (pooling for sex) among lineages, most of the variation in SSD is independent of sex-averaged body size and shows a strong genetic correlation with sex-specific plasticity, such that increased female-biased SSD is associated with increased body size plasticity in females. Our data are consistent with the condition dependence hypothesis of sexual dimorphism and suggest that SSD in Drosophila is a consequence of selection on the developmental genetic mechanisms that regulate the plasticity of body size.

Sexual size and shape dimorphism, and allometric scaling in the pupal and adult traits of Eristalis tenax.

The patterns and amount of variation in size, shape, and/or life history traits between females and males are fundamentally important to gain the comprehensive understanding of the evolution of phenotypic diversity. In addition, the covariation of phenotypic traits can significantly contribute to morphological diversification and sexual dimorphism (SD). Using linear and geometric morphometrics, 237 Eristalis tenax specimens sampled from five populations were, therefore, comparatively assessed for the variation in sexual size dimorphism (SSD), sexual shape dimorphism (SShD), and life history traits, as well as for trait covariation (ontogenetic and static allometry). Pupal body, adult wing, and body mass traits were analyzed. Female-biased SSD was observed for pupal length, width, and centroid size, adult wing centroid size, mass, wing loading, and wing area. Conversely, pupal length/width ratio, developmental time, and mass were not found to be sexually dimorphic. Next, wing SShD, but not pupal body SShD was revealed, while allometry was found to be an important "determinant of SD" at the adult stage, with only a minor impact at the pupal stage. By comparing the patterns of covariance (based on allometric slope and intercept) between respective body mass and morphometric traits of pupae and adults, greater variation in allometric slopes was found in adult traits, while static allometries of the two stages significantly differed, as well. Finally, the results indicate that changes in the allometric intercept could be an important source of intraspecific variation and SD in drone fly adults.

Climate shapes patterns of sexual size and shape dimorphism across the native range of the green anole lizard,Anolis carolinensis(Squamata: Dactyloidae)

AbstractGeographical variation in sexual size dimorphism (SSD) can result from the combined effects of environmental and sexual selection. To understand the determinants of SSD across geographical landscapes, we tested for relationships between SSD and climatic variables in the widespread lizard Anolis carolinensis. To distinguish alternative hypotheses for observed patterns of variation in SSD, we also examined sex-specific patterns of body size evolution and asked whether SSD was associated with certain patterns of sexual shape dimorphism. We found strong evidence for Rensch’s rule (an increase in male-biased SSD with average body size) in A. carolinensis and evidence for the reversed version of Bergmann’s rule (an increase in body size towards warmer environments) in males. Across populations, SSD was positively related to temperature; however, female body size was not related to any climatic variable, suggesting that the latitudinal gradient of SSD might be driven by a gradient in the intensity of sexual selection acting on males. Sexual size dimorphism was positively correlated with sexual dimorphism in head shape and negatively correlated with limb length dimorphism, suggesting that sexual selection in males might drive the evolution of SSD and that differences in size and limb shape between sexes might represent alternative strategies to avoid competition for the same resources.

Sexual size dimorphism and male reproductive traits vary across populations of a tropical rainforest dung beetle species (Onthophagus babirussa).

Sexual size dimorphism (SSD) arises when natural selection and sexual selection act differently on males and females. Male‐biased SSD is rarer in insects and usually indicates strong sexual selection pressure on male body size in a species. Patterns of SSD can also vary between populations of species that are exposed to different environmental conditions, such as differing resource availability and diversity. Here, we investigate intraspecific variation in SSD as well as relative investment in precopulatory (horn length) and postcopulatory traits (sperm length and testes weight) in a tropical rainforest dung beetle Onthophagus babirussa across Singapore and Peninsular Malaysia. Overall, three out of four populations displayed significant male‐biased SSD, and SSD was greater in populations with smaller overall body size. Average male body size was similar across all populations while female body size was significantly smaller in Singapore, suggesting that the pronounced SSD may also be due to stronger sexual selection on male body size in Singapore populations. All populations showed significant investment in horns as a weapon likely used in male‐male competition, while postcopulatory traits showed no clear scaling relationship with body size, suggesting a higher priority on precopulatory sexual traits in the mating system of this species.

Evolution of sexual dimorphism in the plateau brown frog fails to obey Rensch’s rule

Rensch’s rule describes sexual size dimorphism (SSD) that decreases with increasing body size when females are larger than males and SSD that increases when males are larger than females. The plateau brown frog Rana kukunoris, a species endemic to the eastern Tibetan Plateau, exhibits female-biased size dimorphism. Using data on body size from 26 populations and age from 21 populations, we demonstrated that SSD did not increase with increasing mean female snout-vent length (SVL) when controlling for sex-specific age structure, failing to support the Rensch’s rule. Thus, we suggest that fecundity selection (favouring large female size) balances out sexual selection (favouring large male size), which results in a similar divergence between males and females body size. In addition, sex-specific age differences explained most of the variation of SSD across populations.

Evolution of sexual dimorphism in an endemic toad of the Qinghai–Tibet plateau fails to obey Rensch’s rule

According to Rensch's rule, sexual size dimorphism (SSD) decreases with increasing body size in species where females are larger than males, whereas SSD increases with body size when males are larger than females. In this study, we examined body size from 15 populations of Bufo minshanicus in the Qinghai–Tibet plateau in China. We estimated the age of the individuals from eight populations to evaluate Rensch's rule and possible causes of variation in SSD. After or before correcting male and female body size for age differences, the patterns of SSD failed to obey Rensch's rule or its inverse, even though fecundity selection in toads tends to favor larger female body size. We also found that the degree of SSD was positively correlated with the operational sex ratio in the field across B. minshanicus populations. Thus, sexual and fecundity selection play an equal role driving the evolution of SSD within toads. In addition, sex-specific growth rate and age structure explain part of the variation observed in SSD across populations.

Environmental drivers of sexual dimorphism in a lizard with alternative mating strategies.

Understanding the relative importance of sexual and natural selection in shaping morphological traits is a long-standing goal of evolutionary ecology. Male-biased sexual size dimorphism (SSD) is typically associated with male-male competition. Similarly, male polymorphisms are considered a consequence of competitive social interactions. This classic paradigm overlooks the fact that environmental factors mediate social interactions and can lead to ecological adaptations. Common side-blotched lizards, Uta stansburiana, are a model system for this paradigm due to well-known rock-paper-scissors social dynamics between male morphs. SSD in this species has been considered primarily a consequence of social interactions, with male size resulting from the number of morphs in each population and female size being constrained through fecundity benefits. We test if the environment explains intraspecific variation in SSD and number of male morphs in U.stansburiana. By compiling data from 49 populations, we show that environmental variables are stronger predictors of SSD than the number of male morphs. Similarly, we show that the environment mediates SSD and potentially contributes to morph loss in colder environments. We propose that the environment favours smaller males in areas of high seasonality. Our results demonstrate the importance of the environment as a mediator of SSD.

Sex-Specific Population Differences in Resting Metabolism Are Associated with Intraspecific Variation in Sexual Size Dimorphism of Brown Anoles.

AbstractSexual size dimorphism can vary in direction and magnitude across populations, but the extent to which such intraspecific variation is associated with sex and population differences in underlying metabolic processes is unclear. We compared resting metabolic rates (RMRs) of brown anole lizards (Anolis sagrei) from two island populations in the Bahamas (Eleuthera and Great Exuma) that differ in the magnitude of male-biased sexual size dimorphism. Whereas females from each population exhibit similar growth rates and body sizes, males from Great Exuma grow more quickly and attain larger body sizes than males from Eleuthera. We found that these population differences in growth of males persisted in captivity. Therefore, we predicted that males from each population would differ in RMR, whereas females would not. Consistent with this prediction, we found that RMR of males from Eleuthera was higher than that of males from Great Exuma, particularly at higher temperatures. As predicted, RMR of females did not differ between populations. Despite this apparent sex-specific trade-off between growth rate and RMR at the population level, we found a positive relationship between growth rate and RMR at the individual level. The fact that Great Exuma males maintain lower RMR than Eleuthera males, despite their greater absolute growth rates and the positive relationship between RMR and growth rate across individuals, suggests that Great Exuma males may have lower baseline metabolic demands and/or greater growth efficiency than Eleuthera males. Our results call attention to sex-specific divergence in metabolism as a potential mechanism for intraspecific divergence in sexual size dimorphism.

Sexual size dimorphism variation in altitude gradients in ground beetles (Coleoptera, Carabidae)

In this study, we verified the theory about the greatest steepness in body size variation in insects males compared with females in altitude gradients. As the model, ground beetles took as the appreciated bioindicators. We sampled three species of carabids at two regions: Carabus odoratus and Pterostihusmontanus were sampled at four elevations at Barguzin Ridge (N 54° 20’; E 109° 30’, Russia) and Carabus exaratus – at three elevations in mountainous terrain in Chechnya Republic (42088’N 46044’E Russia). We measured more than 1000 individuals by six linear traits and construct regression models of females’ and males’ size variation along elevation gradients. The regression slopes in the majority of the overwhelming case (14 models out of 18) were sex-biased. In P. montanus elytra length variation and C. exaratus traits width variation, the slopes differed in males and females only: In the first case, trait value in females significantly increased and in the second traits values varied in opposite directions. Therefore, we did not find evidence of greater steepness in male traits variation in ground beetles.

Egg-laying site, fecundity and degree of sexual size dimorphism in frogs

AbstractFemale fecundity is an important selective force leading to female-biased sexual size dimorphism (SSD) in frogs. Because anurans exhibit diverse reproductive modes, we investigated whether variation in SSD and fecundity are related with oviposition site. We asked whether arboreal breeding species show pronounced female-biased SSD and if, paradoxically, females have lower fecundity because of the costs of carrying oocytes and amplectant males. Conversely, we tested whether species that deposit eggs in concealed sites show less pronounced SSD, because females do not carry males and space limitation may reduce female size and fecundity. Our results showed that, in general, males were approximately 20% smaller than females. However, for species with hidden oviposition sites, males and females exhibited more similar body sizes and arboreal hylids showed more pronounced female-biased SSD. Overall, fecundity was higher in aquatic breeders, as expected, but in hylids, fecundity was smaller in arboreal breeders, which suggests that arboreality may impose restrictions on fecundity. By analysing SSD in a broader and more specific lineage (Hylidae), we found that reproductive microhabitat may also influence female size and fecundity, playing an important role in the evolution of SSD in frogs at different evolutionary scales.

Altitudinal variation of sexual size dimorphism in ground beetle Carabus odoratus Shill.

Altitudinal variation of sexual size dimorphism in ground beetle Carabus odoratus Shill.

Altitudinal variation of sexual size dimorphism in ground beetle Carabus odoratus Shill.

Samples of Carabus odoratus barguzinicus Shil. сollected in different altitudes of Barguzin Mountain Range at the northeast part of Baikal Lake (N 54° 20’; E 109° 30’, Russia). In total eight populations were analyzed, two in each belt: Coast, Low-, Middle- and High Mountains (455-460, 500-700, 701-1300, 1301-1700 m above sea level). Morphometric measurements included six traits (elytra length and width, pronotum length and width, head length, and the distance between eyes) of 1300 specimens. Sexual size dimorphism (SSD) in Carabus odoratus was female-biased. Its value was significant and more pronounced in Low- and High mountains and varied from 0.01 to 0.1 in different traits and at different altitudes. The largest values of SSD was at Low- and High mountains. The latter considered the most favorable biotopes to the species studied. RMA II method resulted in positive Intercepts in most traits treated. That demonstrated the higher sensitivity of females to environmental conditions and that Carabus odoratus follows the opposite Rensch′s rule.

Male-biased sexual size dimorphism in sharks: the narrowmouth catshark Schroederichthys bivius as case study

In shark populations, variation in sexual size dimorphism (SSD) appears to be related with the reproductive mode. Here, we aimed to investigate the reproductive biology of the narrowmouth catshark Schroederichthys bivius, an oviparous shark with size and secondary sexual dimorphism. For this purpose, specimens were collected during research surveys covering almost their entire distribution in the southwestern Atlantic continental shelf. Size distribution ranged from 196 to 810 mm total length (TL) in males and from 258 to 630 mm TL in females, and the mean size at maturity was 563 mm TL for males and 485 mm TL for females. Males appeared to be capable of mating all times of the year, while females could exhibit an annual reproductive cycle. Besides SSD, with males attaining higher maximum TL and size at maturity, spatial sexual segregation appears to be observed in mature specimens. Although differences in size at maturity were observed between surveys, these differences did not show a latitudinal pattern. The spatial overlap observed between closed areas to trawling and the distribution of egg-bearing females suggests that these closed areas, although designed and established to protect other species, may represent an important measure also for the conservation of S. bivius.

Selection for increased male size predicts variation in sexual size dimorphism among fish species.

Variation in the degree of sexual size dimorphism (SSD) among taxa is generally considered to arise from differences in the relative intensity of male-male competition and fecundity selection. One might predict, therefore, that SSD will vary systematically with (1) the intensity of sexual selection for increased male size, and (2) the intensity of fecundity selection for increased female size. To test these two fundamental hypotheses, we conducted a phylogenetic comparative analysis of SSD in fish. Specifically, using records of body length at first sexual maturity from FishBase, we quantified variation in the magnitude and direction of SSD in more than 600 diverse freshwater and marine fish species, from sticklebacks to sharks. Although female-biased SSD was common, and thought to be driven primarily by fecundity selection, variation in SSD was not dependent on either the allometric scaling of reproductive energy output or fecundity in female fish. Instead, systematic patterns based on habitat and life-history characteristics associated with varying degrees of male-male competition and paternal care strongly suggest that adaptive variation in SSD is driven by the intensity of sexual selection for increased male size.

Intraspecific mating system evolution and its effect on complex male secondary sexual traits: Does male-male competition increase selection on size or shape?

Sexual selection is generally held responsible for the exceptional diversity in secondary sexual traits in animals. Mating system evolution is therefore expected to profoundly affect the covariation between secondary sexual traits and mating success. Whereas there is such evidence at the interspecific level, data within species remain scarce. We here investigate sexual selection acting on the exaggerated male fore femur and the male wing in the common and widespread dung flies Sepsis punctum and S.neocynipsea (Diptera: Sepsidae). Both species exhibit intraspecific differences in mating systems and variation in sexual size dimorphism (SSD) across continents that correlates with the extent of male-male competition. We predicted that populations subject to increased male-male competition will experience stronger directional selection on the sexually dimorphic male foreleg. Our results suggest that fore femur size, width and shape were indeed positively associated with mating success in populations with male-biased SSD in both species, which was not evident in conspecific populations with female-biased SSD. However, this was also the case for wing size and shape, a trait often assumed to be primarily under natural selection. After correcting for selection on overall body size by accounting for allometric scaling, we found little evidence for independent selection on any of these size or shape traits in legs or wings, irrespective of the mating system. Sexual dimorphism and (foreleg) trait exaggeration is therefore unlikely to be driven by direct precopulatory sexual selection, but more so by selection on overall size or possibly selection on allometric scaling.