Patterns of sexual size dimorphism in Chelonia

Macroevolutionary patterns of sexual size dimorphism (SSD) indicate how sexual selection, natural selection, and genetic and developmental constraints mold sex differences in body size. One putative pattern, known as Rensch's rule, posits that, among species with female‐larger SSD, the relative degree of SSD declines with species' body size, whereas, among male‐larger SSD species, relative SSD increases with size. Using a dataset of 196 chelonian species from all fourteen families, we investigated the correlation in body size evolution between male and female Chelonia and the validity of Rensch's rule for the taxon and within its major clades. We conclude that male–female correlations in body size evolution are high, although these correlations differ among chelonian families. Overall, SSD scales isometrically with body size; Rensch's rule is valid for only one family, Testudinidae (tortoises). Because macroevolutionary patterns of SSD can vary markedly among clades, even in a taxon as morphologically conservative as Testudines, one must guard against inappropriately pooling clades in comparative studies of SSD. The results of the present study also indicate that regression models that assume the x‐variable (e.g. male body size) is measured without statistical error, although frequently reported, will result in erroneous conclusions about phylogenetic trends in sexual size dimorphism. © 2012 The Linnean Society of London, Biological Journal of the Linnean Society, 2012, 108 , 396–413.     

Intraspecific variation in body size and sexual size dimorphism,and a test of Rensch's rule in bats

The magnitude and direction of sexual size dimorphism (SSD) may vary considerably within and among taxa, and the primary causes of such variation have not been thoroughly elucidated. For example, the effect of abiotic factors is frequently attributed to explain intra‐ and interspecific variation in SSD. Rensch's rule, which states that males vary more in size than females when body size increases, has rarely been tested in bats. Therefore, whether bats follow Rensch's rule remains unclear, particularly when females are larger than males. We investigated whether four bat species presented SSD, as well as whether their body sizes varied within each sex across localities, testing the hypothesis that intraspecific SSD varies substantially depending of sampling localities. We finally examined whether bats followed Rensch's rule by simultaneously using intraspecific and interspecific approaches. Although SSD was not observed for most bat species within each locality, the females of three of the four captured species exhibited differences in body size between particular localities. Usually the females varied more in size than did males across localities, mostly exhibiting a female‐biased SSD. Significant differences in SSD were observed (i.e. mean values of the sexual dimorphism index), even though Rensch's rule was not followed.     

Lack of support for Rensch's rule in an intraspecific test using red flour beetle (Tribolium castaneum) populations

Rensch's rule proposes a universal allometric scaling phenomenon across species where sexual size dimorphism (SSD) has evolved: in taxa with male‐biased dimorphism, degree of SSD should increase with overall body size, and in taxa with female‐biased dimorphism, degree of SSD should decrease with increasing average body size. Rensch's rule appears to hold widely across taxa where SSD is male‐biased, but not consistently when SSD is female‐biased. Furthermore, studies addressing this question within species are rare, so it remains unclear whether this rule applies at the intraspecific level. We assess body size and SSD within Tribolium castaneum (Herbst), a species where females are larger than males, using 21 populations derived from separate locations across the world, and maintained in isolated laboratory culture for at least 20 years. Body size, and hence SSD patterns, are highly susceptible to variations in temperature, diet quality and other environmental factors. Crucially, here we nullify interference of such confounds as all populations were maintained under identical conditions (similar densities, standard diet and exposed to identical temperature, relative humidity and photoperiod). We measured thirty beetles of each sex for all populations, and found body size variation across populations, and (as expected) female‐biased SSD in all populations. We test whether Rensch's rule holds for our populations, but find isometry, i.e. no allometry for SSD. Our results thus show that Rensch's rule does not hold across populations within this species. Our intraspecific test matches previous interspecific studies showing that Rensch's rule fails in species with female‐biased SSD.     

Sexual dichromatism in wing pigmentation of New World dragonflies follows Rensch's rule

Many animal taxa that display sexual size dimorphism (SSD) exhibit a positive allometric relationship in which the degree of dimorphism increases with body size. This macroevolutionary pattern is known as Rensch's rule. Although sexual selection is hypothesized to be the main mechanism causing this pattern, body size is influenced by several selective forces, including natural and sexual selection. Therefore, by focusing exclusively on SSD one cannot ascertain which of these selective forces drives Rensch's rule. If sexual selection is indeed the main mechanism underlying Rensch's rule, we predict that other sexually selected traits, including coloration‐based ornaments, will also exhibit interspecific allometric scaling consistent with Rensch's rule. We tested this prediction using wing pigmentation of 89 species of dragonflies. Studies show that male wing pigmentation is generally under strong intra‐ and intersexual selection, so that sexual dichromatism in this trait should follow Rensch's rule. Conversely, the available evidence suggests that male body size is usually not sexually selected in dragonflies, so we do not expect SSD to follow Rensch's rule. First, we found that sexual dichromatism in wing pigmentation was consistent with Rensch's rule. The phylogenetic major axis regression slope was significantly greater than one. We also showed that the allometric slope for SSD was not different from unity, providing no support for Rensch's rule. Our results provide the first evidence that a trait which appears to be under strong sexual selection exhibits a pattern consistent with Rensch's rule.     

Function‐related Drivers of Skull Morphometric Variation and Sexual Size Dimorphism in a Subterranean Rodent,Plateau Zokor (Eospalax baileyi)

Sexual dimorphism is prevalent in most living organisms. The difference in size between sexes of a given species is generally known as sexual size dimorphism (SSD). The magnitude of the SSD is determined by Rensch's rule where size dimorphism increases with increasing body size when the male is the larger sex and decreases with increasing average body size when the female is the larger sex. The unique underground environment that zokors (Eospalax baileyi) live under in the severe habitat of the Qinghai‐Tibetan Plateau (QTP) could create SSD selection pressures that may or may not be supported by Rensch's rule, making this scientific question worthy of investigation. In this study, we investigated the individual variation between sexes in body size and SSD of plateau zokors using measurements of 19 morphological traits. We also investigated the evolutionary mechanisms underlying SSD in plateau zokors. Moreover, we applied Rensch's rule to all extant zokor species. Our results showed male‐biased SSD in plateau zokors: The body‐ and head‐related measurements were greater in males than in females. Linear regression analysis between body length, body weight, and carcass weight showed significant relationships with some traits such as skull length, lower incisor length, and tympanic bulla width, which might support our prediction that males have faster growth rates than females. Further, the SSD pattern corroborated the assumption of Rensch's rule in plateau zokors but not in the other zokor species. Our findings suggest that the natural underground habitat and behavioral differences between sexes can generate selection pressures on male traits and contribute to the evolution of SSD in plateau zokors.     

A test of Rensch's rule in varanid lizards

In a model group of giant reptiles, we explored the allometric relationships between male and female body size and compared the effects of sexual and fecundity selection, as well as some proximate causes, on macroevolutionary patterns of sexual size dimorphism (SSD). Monitor lizards are a morphologically homogeneous group that has been affected by extreme changes in body size during their evolutionary history, resulting in 14‐fold differences among the body sizes of recent species. Here, we analysed data concerning the maximum and/or mean male and female snout–vent lengths in 42 species of monitor lizard from literary sources and supplemented these data with measurements made in zoos. There was a wide scale of SSD from nearly monomorphic species belonging mostly to the subgenus Odatria and Prasinus group of the Euprepriosaurus to apparently male‐larger taxa. The variable best explaining SSD was the body size itself; the larger the species, the higher the SSD. This pattern agrees with the currently discussed Rensch's rule, claiming that the relationship between male and female body size is hyperallometric, i.e. the allometric exponent of this relationship exceeds unity and thus SSD increases with body size in the case of male‐larger taxa. All our estimates of the reduced major axis regression slopes of this relationship ranged from 1.132 to 1.155. These estimates are significantly higher than unity, and thus unequivocally corroborate the validity of Rensch's rule in this reptilian group. In spite of our expectation that the variation in SSD can be alternatively explained by variables reflecting the strength of sexual selection (presence of male combat), fecundity selection (e.g. clutch size and mass) and/or proximate ecological factors (habitat type), none of these variables had consistent effects on SSD, especially when the data were adjusted to phylogenetic dependence and/or body size. © 2010 The Linnean Society of London, Biological Journal of the Linnean Society, 2010, 100 , 293–306.     

Sexual size dimorphism in musteloids: An anomalous allometric pattern is explained by feeding ecology

Rensch's rule states that sexual size dimorphism (SSD) increases with body size in taxa where males are larger, and decreases when females are larger. The dominant explanation for the trend is currently that competitive advantage for males is greater in larger individuals, whereas female size is constrained by the energetics of rearing offspring. This rule holds for a variety of vertebrate taxa, and opposing trends are rare. We examine the allometry of SSD within the Musteloidea and demonstrate a hypo‐allometry contrary to Rensch's rule, with lower SSD associated with larger body size. We provide evidence that feeding ecology is involved. Where diet promotes group‐living, the optimal strategy for the males of larger species is often not to attempt to defend access to multiple females, obviating any competitive advantage of relatively greater size. We conclude that the effect of feeding ecology on mating systems may be a hitherto neglected factor explaining variation in SSD.     

Sexual selection explains sex-specific growth plasticity and positive allometry for sexual size dimorphism in a reef fish

In 1950, Rensch noted that in clades where males are the larger sex, sexual size dimorphism (SSD) tends to be more pronounced in larger species. This fundamental allometric relationship is now known as ‘Rensch''s rule’. While most researchers attribute Rensch''s rule to sexual selection for male size, experimental evidence is lacking. Here, we suggest that ultimate hypotheses for Rensch''s rule should also apply to groups of individuals and that individual trait plasticity can be used to test those hypotheses experimentally. Specifically, we show that in the sex-changing fish Parapercis cylindrica, larger males have larger harems with larger females, and that SSD increases with harem size. Thus, sexual selection for male body size is the ultimate cause of sexual size allometry. In addition, we experimentally illustrate a positive relationship between polygyny potential and individual growth rate during sex change from female to male. Thus, sexual selection is the ultimate cause of variation in growth rate, and variation in growth rate is the proximate cause of sexual size allometry. Taken together, our results provide compelling evidence in support of the sexual selection hypothesis for Rensch''s rule and highlight the potential importance of individual growth modification in the shaping of morphological patterns in Nature.     

Patterns of sexual size dimorphism in Chelonia: revisiting Kinosternidae

Rensch's rule, a macroevolutionary pattern in which sexual size dimorphism (SSD) increases with body size in male‐biased SSD species, or decreases with female‐biased SSD species, has been investigated in many vertebrates because it indicates whether SSD is being driven by sexual selection or a different force (i.e. fecundity or natural selection). Evidence in turtles has shown some conflicting results, which may be explained by the different phylogenies used in the analyses. Because the newly available well‐resolved phylogeny of family Kinosternidae provides evidence for the ancient monophyly of Staurotypidae and Kinosternidae and their recognition as separate families (previously Staurotypidae was considered as a subfamily within Kinosternidae) and introduced the genus Cryptochelys for the monophyletic leucostomum clade, we revisit the pattern of SSD and body size in Kinosternidae. By contrast to what had been proposed, we found that the Kinosternidae as formerly recognized (i.e. including Staurotypus and Claudius) and the restricted Kinosternidae both follow a pattern consistent with Rensch's rule. Our analysis with published body size data did not change our results, confirming the importance of the phylogeny used in macroevolutionary studies. © 2014 The Linnean Society of London, Biological Journal of the Linnean Society, 2014, 111 , 806–809.     

Patterns of sexual size dimorphism in stingless bees: Testing Rensch’s rule and potential causes in highly eusocial bees (Hymenoptera: Apidae,Meliponini)

Eusocial insects offer a unique opportunity to analyze the evolution of body size differences between sexes in relation to social environment. The workers, being sterile females, are not subject to selection for reproductive function providing a natural control for parsing the effects of selection on reproductive function (i.e., sexual and fecundity selection) from other kinds of natural selection. Patterns of sexual size dimorphism (SSD) and testing of Rensch's rule controlling for phylogenetic effects were analyzed in the Meliponini or stingless bees. Theory predicts that queens may exhibit higher selection for fecundity in eusocial taxa, but contrary to this, we found mixed patterns of SSD in Meliponini. Non‐Melipona species generally have a female‐biased SSD, while all analyzed species of Melipona showed a male‐biased SSD, indicating that the direction and magnitude of the selective pressures do not operate in the same way for all members of this taxon. The phylogenetic regressions revealed that the rate of divergence has not differed between the two castes of females and the males, that is, stingless bees do not seem to follow Rensch's rule (a slope >1), adding this highly eusocial taxon to the various solitary insect taxa not conforming with it. Noteworthy, when Melipona was removed from the analysis, the phylogenetic regressions for the thorax width of males on queens had a slope significantly smaller than 1, suggesting that the evolutionary divergence has been larger in queens than males, and could be explained by stronger selection on female fecundity only in non‐Melipona species. Our results in the stingless bees question the classical explanation of female‐biased SSD via fecundity and provide a first evidence of a more complex determination of SSD in highly eusocial species. We suggest that in highly eusocial taxa, additional selection mechanisms, possibly related to individual and colonial interests, could influence the evolution of environmentally determined traits such as body size.     

Sexual size dimorphism in domestic goats,sheep, and their wild relatives

Sexual size dimorphism (SSD) is a widespread phenomenon in different animal taxa, including the subfamily of goats and sheep (Caprinae), which belongs to the most dimorphic mammalian groups. Rensch's rule describes the pattern of SSD, claiming that larger species generally exhibits higher male to female body size ratio. Agreement with Rensch's rule is manifested by slope of the allometric relationship between male and female body size exceeding one. To test this rule, we analysed the data available in the literature on adult body mass of males and females in domestic goat and sheep breeds (169 and 303, respectively) and 37 wild species/subspecies of the subfamily Caprinae. According to the current phylogenetical hypotheses, there are six distinct monophyletic groups with different levels of SSD (expressed as M/F): (1) wild goats (1.83); (2) wild sheep (1.67); (3) non‐European chamoises, including Ovibos moschatus (1.18); (4) European chamoises (1.27); (5) Budorcas taxicolor (1.01); and (6) Pantholops hodgsonii (1.65). Domestication has led to a remarkable decline in SSD of both domestic goats (1.36) and sheep (1.41). The highest regression slope of the relationship between male and female body size is that estimated for wild goats (1.32), followed by wild sheep (1.24), non‐European chamoises (1.14), domestic sheep (1.13), and domestic goats (1.10). Nevertheless, only the last two values are statistically different from one and thus corroborate Rensch's rule. © 2009 The Linnean Society of London, Biological Journal of the Linnean Society, 2009, 98 , 872–883.     

The evolution of sexual size dimorphism in cottontail rabbits (Sylvilagus,Leporidae)

In mammals, ‘female‐biased’ sexual size dimorphism (SSD), in which females are larger than males, is uncommon. In the present study, we examined Sylvilagus, a purported case of female‐biased SSD, for evolutionary correlations among species between SSD, body‐size, and life‐history variables. We find that: (1) although most species are female‐biased, the degree and direction of SSD vary more than was previously recognized and (2) the degree of SSD decreases with increasing body size. Hence, Sylvilagus provides a new example, unusual for a female‐biased taxon, in which allometry for SSD is consistent with ‘Rensch's Rule’. As a corollary to Rensch's Rule, we observe that changes in SSD in Sylvilagus are typically associated with larger, more significant changes in males than females. Female‐biased SSD could be produced by selection for larger females, smaller males, or both. Although larger female size may be related to high fecundity and the extremely rapid fetal and neonatal growth in Sylvilagus, we find little evidence for a correlation between SSD and various fecundity‐related traits in among‐species comparisons. Smaller male size may confer greater reproductive success through greater mobility and reduced energetic requirements. We propose that a suite of traits (female dispersion, large male home ranges, reduced aggression, and a promiscuous mating system) has favoured smaller males and thus influenced the evolution of SSD in cottontails. © 2008 The Linnean Society of London, Biological Journal of the Linnean Society, 2008, 95 , 141–156.     

Equal temperature–size responses of the sexes are widespread within arthropod species

Sexual size dimorphism (SSD) is often affected by environmental conditions, but the effect of temperature on SSD in ectotherms still requires rigorous investigation. We compared the plastic responses of size-at-maturity to temperature between males and females within 85 diverse arthropod species, in which individuals of both sexes were reared through ontogeny under identical conditions with excess food. We find that the sexes show similar relative (proportional) temperature–body size (T–S) responses on average. The high degree of similarity occurs despite an analysis that includes a wide range of animal body sizes, variation in degree of SSD and differences in the sign of the T–S response. We find no support for Rensch''s rule, which predicts greater variation in male size, or indeed the reverse, greater female size variation. SSD shows no systematic temperature dependence in any of the 17 arthropod orders examined, five of which (Diptera, Orthoptera, Lepidoptera, Coleoptera and Calanoida) include more than six thermal responses. We suggest that the same proportional T–S response may generally have equivalent fitness costs and benefits in both sexes. This contrasts with effects of juvenile density, and food quantity/quality, which commonly result in greater size plasticity in females, suggesting these variables have different adaptive effects on SSD.     

Allometry of Sexual Size Dimorphism in Domestic Dog

Background

The tendency for male-larger sexual size dimorphism (SSD) to scale with body size – a pattern termed Rensch''s rule – has been empirically supported in many animal lineages. Nevertheless, its theoretical elucidation is a subject of debate. Here, we exploited the extreme morphological variability of domestic dog (Canis familiaris) to gain insights into evolutionary causes of this rule.

Methodology/Principal Findings

We studied SSD and its allometry among 74 breeds ranging in height from less than 19 cm in Chihuahua to about 84 cm in Irish wolfhound. In total, the dataset included 6,221 individuals. We demonstrate that most dog breeds are male-larger, and SSD in large breeds is comparable to SSD of their wolf ancestor. Among breeds, SSD becomes smaller with decreasing body size. The smallest breeds are nearly monomorphic.

Conclusions/Significance

SSD among dog breeds follows the pattern consistent with Rensch''s rule. The variability of body size and corresponding changes in SSD among breeds of a domestic animal shaped by artificial selection can help to better understand processes leading to emergence of Rensch''s rule.     

Does ‘gliding while gravid’ explain Rensch's rule in flying lizards?

Among species with sexual size dimorphism (SSD), taxa in which males are the larger sex have increasing SSD with increasing body size, whereas in taxa in which females are the larger sex, SSD decreases with body size: Rensch's rule. We show in flying lizards, a clade of mostly female‐larger species, that SSD increases with body size, a pattern similar to that in clades with male‐biased SSD or more evenly mixed SSD. The observed pattern in Draco appears due to SSD increasing with evolutionary changes in male body size; specifically divergence in body size among species that are in sympatric congeneric assemblages. We suggest that increasing body size, resulting in decreased gliding performance, reduces the relative gliding cost of gravidity in females, and switches sexual selection in males away from a small‐male, gliding advantage and toward selection on large size and fighting ability as seen in many other lizards. Thus, selection for large females is likely greater than selection for large males at the smaller end of the body size continuum, whereas this relationship reverses for species at the larger end of the continuum. © 2014 The Linnean Society of London, Biological Journal of the Linnean Society, 2014, 113 , 270–282.     

Body size and elevation: do Bergmann's and Rensch's rule apply in the polytypic bushcricket Poecilimon veluchianus?

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Ecological drivers of body size evolution and sexual size dimorphism in short‐horned grasshoppers (Orthoptera: Acrididae)

Sexual size dimorphism (SSD) is widespread and variable in nature. Although female‐biased SSD predominates among insects, the proximate ecological and evolutionary factors promoting this phenomenon remain largely unstudied. Here, we employ modern phylogenetic comparative methods on eight subfamilies of Iberian grasshoppers (85 species) to examine the validity of different models of evolution of body size and SSD and explore how they are shaped by a suite of ecological variables (habitat specialization, substrate use, altitude) and/or constrained by different evolutionary pressures (female fecundity, strength of sexual selection, length of the breeding season). Body size disparity primarily accumulated late in the history of the group and did not follow a Brownian motion pattern, indicating the existence of directional evolution for this trait. We found support for the converse of Rensch's rule (i.e. females are proportionally bigger than males in large species) across all taxa but not within the two most speciose subfamilies (Gomphocerinae and Oedipodinae), which showed an isometric pattern. Our results do not provide support for the fecundity or sexual selection hypotheses, and we did not find evidence for significant effects of habitat use. Contrary to that expected, we found that species with narrower reproductive window are less dimorphic in size than those that exhibit a longer breeding cycle, suggesting that male protandry cannot solely account for the evolution of female‐biased SSD in Orthoptera. Our study highlights the need to consider alternatives to the classical evolutionary hypotheses when trying to explain why in certain insect groups males remain small.     

A global assessment of Bergmann's rule in mammals and birds

Bergmann's rule states that endotherms have a large body size in high latitudes and cold climates. However, previous empirical studies have reported mixed evidence on the relationships between body size and latitude, raising the question of why some clades of endotherms follow Bergmann's rule, whereas others do not. Here, we synthesized the interspecific relationships between body size and latitude among 16,187 endothermic species (5422 mammals and 10,765 birds) using Bayesian phylogenetic generalized linear mixed models to examine the strength and magnitude of Bergmann's rule. We further assessed the effect of biological and ecological factors (i.e., body mass categories, dietary guild, winter activity, habitat openness, and climate zone) on the variations in the body mass–latitude relationships by adding an interaction term in the models. Our results revealed a generally weak but significant adherence to Bergmann's rule among all endotherms at the global scale. Despite taxonomic variation in the strength of Bergmann's rule, the body mass of species within most animal orders showed an increasing trend toward high latitudes. Generally, large-bodied, temperate species, non-hibernating mammals, and migratory and open-habitat birds tend to conform to Bergmann's rule more than their relatives do. Our results suggest that whether Bergmann's rule applies to a particular taxon is mediated by not only geographic and biological features, but also potential alternate strategies that species might have for thermoregulation. Future studies could explore the potential of integrating comprehensive trait data into phylogenetic comparative analysis to re-assess the classic ecogeographic rules on a global scale.     

Life history responses of the cabbage beetle Colaphellus bowringi to temperature change

Temperature is considered one of the most important mediators of phenotypic plasticity in ectotherms. Here, we investigated life history traits of the cabbage beetle, Colaphellus bowringi Baly (Coleoptera: Chrysomelidae), at a wide range of temperatures (16, 19, 22, 24, 26 and 28°C). The larval and pupal times were significantly decreased with increasing rearing temperature and growth rate was positively correlated with temperature. However, the relationship between body size and rearing temperature in C. bowringi did not follow the temperature–size rule; both males and females reached the highest body weight at 19°C. Females were significantly larger than males at all temperatures. Male pupae lost significantly more weight at metamorphosis compared to females. However, diapausing males gained significantly higher weight after feeding compared to diapausing females at higher temperatures of 22, 24, 26 and 28°C. Body weight tended to decrease with increasing rearing temperature, whereas sexual size dimorphism (SSD) tended to increase with increasing rearing temperature; thus, Rensch's rule is upheld. The degree to which SSD changed with temperature varied with different development stages. SSD was lowest in pupae, highest in newly emerged adults and intermediate in diapausing adults.     

It is not always about body size: evidence of Rensch's rule in a male weapon

In many species, sexual dimorphism increases with body size when males are the larger sex but decreases when females are the larger sex, a macro-evolutionary pattern known as Rensch''s rule (RR). Although empirical studies usually focus exclusively on body size, Rensch''s original proposal included sexual differences in other traits, such as ornaments and weapons. Here, we used a clade of harvestmen to investigate whether two traits follow RR: body size and length of the fourth pair of legs (legs IV), which are used as weapons in male–male fights. We found that males were slightly smaller than females and body size did not follow RR, whereas legs IV were much longer in males and followed RR. We propose that sexual selection might be stronger on legs IV length than on body size in males, and we discuss the potential role of condition dependence in the emergence of RR.