Sexual Size Dimorphism in Anurans: Ontogenetic Determination Revealed by an Across-Species Comparison

Ever since Darwin, why and how sexual size dimorphism (SSD) arises among the animal kingdom has puzzled evolutionary biologists. While comparative analyses focus on ultimate causes of this phenomenon, few attempts have been made to quantify its ontogenetic basis. Here we deal with the latter issue using the skeletochronology-based demographic data gathered from the literature for anurans, a group of indeterminately growing vertebrates typically exhibit female-biased SSD. Both non- and phylogenetically corrected analyses showed (1) SSD was correlated positively with the sex difference in adult mean age but negatively with the corresponding difference in annual growth rate, (2) the sex difference in adult mean age tended to be negatively correlated with that in growth rate, and (3) the sex difference in adult mean age was positively correlated with that in age at maturity. We conclude that in anuran lineages, the degree of SSD bias towards females is mediated by sexual bimaturation: females mature later than males, and thus live longer and attain larger despite growing slower. Stronger fecundity selection benefits to females, despite incurring growth cost, in combination with the benefits to males from increased reproductive opportunities resulting from earlier maturity, should be the primary evolutionary drivers of the typically female-biased SSD.     

The Proximate Causes of Sexual Size Dimorphism in Phrynocephalus przewalskii

Sexual size dimorphism (SSD) is a common phenomenon and is a central topic in evolutionary biology. Recently, the importance of pursuing an ontogenetic perspective of SSD has been emphasized, to elucidate the proximate physiological mechanisms leading to its evolution. However, such research has seldom focused on the critical periods when males and females diverge. Using mark-recapture data, we investigated the development of SSD, sex-specific survivorship, and growth rates in Phrynocephalus przewalskii (Agamidae). We demonstrated that both male and female lizards are reproductively mature at age 10–11 months (including 5 months hibernation). Male-biased SSD in snout-vent length (SVL) was only found in adults and was fully expressed at age 11 months (June of the first full season of activity), just after sexual maturation. However, male-biased SSD in tail length (TL), hind-limb length (LL), and head width (HW) were fully expressed at age 9–10 months, just before sexual maturation. Analysis of age-specific linear growth rates identified sexually dimorphic growth during the fifth growth month (age 10–11 months) as the proximate cause of SSD in SVL. The males experienced higher mortality than females in the first 2 years and only survived better than females after SSD was well developed. This suggests that the critical period of divergence in the sizes of male and female P. przewalskii occurs between 10 and 11 months of age (May to June during the first full season of activity), and that the sexual difference in growth during this period is the proximate cause. However, the sexual difference in survivorship cannot explain the male-biased SSD in SVL. Our results indicate that performance-related characteristics, such as TL, HW, and LL diverged earlier than SVL. The physiological mechanisms underlying the different growth patterns of males and females may reflect different energy allocations associated with their different reproductive statuses.     

Size and scaling of sexually-selected traits in the lizard, Uta palmeri

Differences between the sexes in overall body size and in the size of other morphological traits, relative to overall body size, are common in many animals. In this study, patterns of growth and scaling of sexually dimorphic tratis are assessedin a lilzard and then used to sugest general developmental mechanisms responsible for sexual size dimorphism (SSD). Adult make Uta palmeri lizards are larger than adult females inoverall body size (snout-vent length, SVL), body mass, jaw length head width, and head depth. Two general growth processes produce this adult SSD. First, juvenile males have greater annual SVL growth rates than do juvenile females, contributing to adult SSD because males will be larger than females in any trait positively correlated with SVL. Secondly, males and females differ in age-related changes in growth of the three head size traits, relative to growth in SVL. Comparing slopes from reduced major axis regressions of each trait on SVL reveals that the sexes do not differ in the scaling of these traits as juveniles, but as adults males have greater slopes than adult females, indicating ontogenetic differences in scaling of these traits in males. Two other topics in SSD are addressed with these data. First, comparing these data on scaling to those of an earlier analysis that used ordinary least squares regression reveals that conclusions about underlying mechanisms in an analysis of scaling can be altered by the choice of a regression model. Secondly, these data indicate that postmaturational differences in scaling contribute to adult sexual size differences, contrary to an earlier study. Shine (1990) found that for many ectotherms, which continue to grow after sexual maturation, post-maturational events contribute little to sexual differences in overall body size. Results for U. palmeri suggest that these findings may only hold for measures of overall body size (e.g. SVL) and may not generalize to traits that exhibit sex difference in scaling.     

Sex Ratio and Sexual Size Dimorphism in a Toad-headed Lizard,Phrynocephalus guinanensis

Phrynocephalus guinanensis has sexual dimorphism in abdominal coloration, but its ontogenetic development of sexual size dimorphism(SSD) is unknown. Using mark-recapture data during four days each year from August from 2014 to 2016, we investigated the development of sex ratios, SSD, sex-specific survivorship and growth rates in a population of P. guinanensis. Our results indicated that the sex ratio of males to females was 1:2.8. Males had a lower survival rate(6%) than females(14%) across the age range from hatchling to adult, which supported the discovered female-biased sex ratio potentially associated with the low survival rate of males between hatchlings and juveniles. Male-biased SSD in tail length and head width existed in adults rather than in hatchling or juvenile lizards. The growth rates in body dimensions were undistinguishable between the sexes during the age from hatchling to juvenile, but the growth rate in head length from juvenile to adult was significantly larger in males than females. Average growth rate of all morphological measurements from hatchling to juvenile were larger compared with corresponding measurements from juvenile to adult, but only being significant in tail length, head width, abdomen length in females and snout-vent length in males. We provided a case study to strengthen our understanding of the important life history traits on how a viviparous lizard population can survive and develop their morphology in cold climates.     

Evolution of Sexual Size Dimorphism in a Frog Obeys the Inverse of Rensch’s Rule

Rensch’s rule refers to a pattern in sexual size dimorphism (SSD) in which SSD increases with body size when males are the larger sex and decreases with body size when females are the larger sex. Using data on body size from 40 populations and age from 31 populations of the rice frog Rana limnochari with female-biased size dimorphism, I tested the consistency of allometric relationships between males and females with Rensch’s rule and evaluated the hypothesis that SSD was largely a function of age differences between the sexes. Statistical comparisons of body sizes between the sexes showed the evidence for the inverse of Rensch’s rule, indicating the level of SSD increased with increasing mean body size. One of the explanations for the occurrence of the inverse of Rensch’s rule may be the fecundity selection hypothesis assuming increased reproductive output in large females. However, differences in age between males and females among populations could explain mildly the variation in SSD.     

Ontogeny of sexual size dimorphism in monitor lizards: males grow for a longer period, but not at a faster rate

Monitor lizards belong to the largest and the most sexually dimorphic lizards in terms of size, making this group an ideal model for studies analyzing ontogenetic causes of sexual dimorphism. Understanding of these ontogenetic factors is essential to the current discussion concerning patterns of sexual dimorphism in animals. We examined the ontogenetic trajectories of body weight and snout-vent length to analyze the emergence of sexual size dimorphism. Experimental animals were 22 males and 13 females of mangrove-dwelling monitors (Varanus indicus) hatched at the Prague Zoo. They were regularly weighed and measured up to the age of 33-40 months, and subsequently sexed by ultrasonographic imaging. The logistic growth equation was used to describe and analyze the observed growth patterns. Our results confirm considerable sexual size dimorphism in the mangrove monitor. The mean asymptotic body weight of males was nearly three times higher than that of females. As the body size of male and female hatchlings is almost equal, and the growth rate parameter (K) of the logistic growth equation as well as the absolute growth rate up to the age of 12 months do not differ between the sexes, size differences between fully grown males and females should be attributed to timing of the postnatal growth. Males continue to grow several months after they reach the age when the growth of females is already reduced. Therefore, the sexual size dimorphism emerges and sharply increases at this period.     

Proximate causes of sexual size dimorphism in Colorado pikeminnow, a long‐lived cyprinid

Evidence for sexual size dimorphism (SSD) and its possible causes were examined in the endangered Colorado pikeminnow Ptychocheilus lucius, a large, piscivorous, cyprinid endemic to the Colorado River system of North America. Individuals representing 18–24% of the upper Colorado River population were captured, measured, sexed and released in 1999 and 2000. Differing male and female total length‐(L_T) frequency distributions revealed SSD with females having greater mean and maximum sizes than males. Although both sexes exhibit indeterminate post‐maturity growth, growth trajectories differed. The point of trajectory divergence was not established, but slowed male growth might coincide with the onset of maturation. Differing growth rate was the dominant proximate cause of SSD, accounting for an estimated 61% of the observed difference in mean adult L_T. The degree of SSD in adults, however, was also related to two other factors. Evidence suggests males become sexually active at a smaller size and earlier age than females; a 2 year difference, suggested here, accounted for an estimated 12% of the between‐sex difference in mean adult L_T. Temporal shifts in gender‐specific survival accounted for an additional 27% of the observed between‐sex difference in mean adult L_T. Estimated age distributions indicated a higher number of older females than older males and more younger males than younger females in the population during the period of sampling. Dissimilarity of age distributions was an unexpected result because the male : female population sex ratio was 1 : 1 and estimates of long‐term annual survival for adult males and females were equal (88%). Future assessments of SSD in this population are apt to vary depending on the prior history of short‐term gender‐specific survival. Without recognizing SSD, non‐gender‐specific growth curves overestimate mean age of adult females and underestimate mean age of adult males of given L_T. Assuming age 8 years for first reproduction in males and age 10 years for females, the adult male : female ratio was estimated as 1·1 : 1 and mean adult age, or generation time, was estimated as 16·4 years for males and 18·4 years for females.     

Proximate developmental mediators of sexual dimorphism in size: case studies from squamate reptiles

Sexual dimorphism in size (sexual size dimorphism; SSD) is nearlyubiquitous, but the relative importance of genetic versus environmentalcontrol of SSD is not known for most species. We investigatedproximate determinants of SSD in several species of squamatereptiles, including three species of Sceloporus lizards andthe diamond-backed rattlesnake (Crotalus atrox). In naturalpopulations of these species, SSD is caused by sexual differencesin age-specific growth. Males and females, however, may oftenshare similar potentials for growth: growth is strongly responsiveto the availability of food, and sexual differences in growthcan be greatly suppressed or completely absent under commonenvironmental conditions in the laboratory. Sexually divergentgrowth is expressed in natural environments because of inherentecological differences between males and females and becauseof potential epigenetic effects of sex-specific growth regulators.In field-active Sceloporus, sexual differences in growth rateare associated with sexual divergence in plasma testosterone.Experiments confirm that testosterone inhibits growth in speciesin which females are larger (for example, S. undulatus and S.virgatus) and stimulates growth in those in which males arelarger (for example, S. jarrovii). Interestingly, however, sexualdivergence in plasma testosterone is not accompanied by divergencein growth in S. jarrovii or in male-larger C. atrox in the laboratory.Furthermore, experimental effects of castration and testosteronereplacement on growth are not evident in captive S. jarrovii,possibly because growth effects of testosterone are supersededby an abundant, high-quality diet. In female-larger S. undulatus,growth may be traded-off against testosterone-induced reproductivecosts of activity. In male-larger species, costs of reproductionin terms of growth are suggested by supplemental feeding ofreproductive female C. atrox in their natural environment andby experimental manipulation of reproductive cost in femaleS. jarrovii. Growth costs of reproduction, however, do not contributesubstantially to the development of SSD in male-larger S. jarrovii.We conclude that the energetic costs of testosterone-induced,male reproductive behavior may contribute substantially to thedevelopment of SSD in some female-larger species. However, despitestrong evidence that reproductive investment exacts a substantialcost in growth, we do not support the reproductive cost hypothesisas a general explanation of SSD in male-larger species.     

Incorporating an ontogenetic perspective into evolutionary theory of sexual size dimorphism

Sexual size dimorphism (SSD) describes divergent body sizes of adult males and females. While SSD has traditionally been explained by sexual and fecundity selection, recent advances in physiology and developmental biology emphasize that SSD would occur proximately because of sexual differences in ontogenetic growth trajectories (i.e., growth rate and duration). Notably, these ontogenetic traits are subject to energetic or time constraints and thus traded off with fitness components (e.g., survival and reproduction). To elucidate the importance of such ontogenetic trade‐offs in the evolution of SSD, we developed a new theoretical framework by extending quantitative genetic models for the evolution of sexual dimorphism in which we reinterpret the trait as body size and reformulate sex‐specific fitness in size‐dependent manners. More specifically, we assume that higher growth rate or longer growth duration leads to larger body size and higher reproductive success but incurs the cost of lower survivorship or shorter reproduction period. We illustrate how two sexes would optimize ontogenetic growth trajectories in sex‐specific ways and exhibit divergent body sizes. The present framework provides new insights into the evolutionary theory of SSD and predictions for empirical testing.     

Experimental and demographic analyses of growth rate and sexual size dimorphism in a lizard, Sceloporus undulatus

Sexual size dimorphism (SSD) is a common phenomenon caused by a variety of environmental and genetic mechanisms in animals. In the current study, we investigate the demography of a population of eastern fence lizards ( Sceloporus undulatus ) to compare age structure and survivorship between the sexes, and we examine growth rates of juveniles under both natural and controlled laboratory conditions to elucidate causes of SSD in this species. Furthermore, using our laboratory growth data, we examine the heritability of juvenile growth rates. Our results show that SSD develops in the field before the end of the first year of age (before sexual maturity) because juvenile females grow more rapidly than juvenile males. In the laboratory environment, however, we observed no sexual difference in growth rates for lizards up to the size of maturity in the field. Thus, sexual differences in growth rate and subsequent development of SSD in this population are highly plastic and subject to strong proximal control. We found high levels of additive genetic variance for juvenile growth, indicating a strong potential for selection to operate on juvenile growth rates. Our results indicate that selection on juvenile growth rate could account for differences in growth among populations but would not necessarily contribute to SSD within our population due to the high plasticity in growth rate.     

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.     

Ontogeny and sexual dimorphism in a captive population of juvenile striped skunks <Emphasis Type="Italic">Mephitis mephitis</Emphasis>

Three main hypotheses can explain the origin of the sexual size dimorphism: (1) the birth-size hypothesis, which states that birth size of males is larger than that of females; (2) the growth-rate hypothesis, which states that males grow faster than females; (3) the growth-length hypothesis, which states that males grow for a longer period of time than females. We examined the factors that may contribute to sexual size dimorphism with growth data of striped skunks Mephitis mephitis Schreber, 1776 held in captivity in Manitoba (Canada), from 7 to 72 days of age. At seven days of age, the mass of male skunks (mean = 79.7 g ± 13.9 SE, n = 37) was significantly larger than that of females (mean = 71.2 g ± 15.0 SE, n = 35) but the head and body length was not statistically different between males (mean = 110.3 mm ± 8.0 SE, n = 37) and females (mean = 95.3 mm ± 7.4 SE, n = 35). There was no difference in growth rate for mass or for length between sexes. We were not able to test for a difference in growth length between sexes. Our results suggest that mass dimorphism occurs early in life.     

THE EVOLUTION OF FEMALE-BIASED SEXUAL SIZE DIMORPHISM: A POPULATION-LEVEL COMPARATIVE STUDY IN HORNED LIZARDS (PHRYNOSOMA)

Female-biased sexual size dimorphism is uncommon among vertebrates and traditionally has been attributed to asymmetric selective pressures favoring large fecund females (the fecundity-advantage hypothesis) and/or small mobile males (the small-male advantage hypothesis). I use a phylogenetically based comparative method to address these hypotheses for the evolution and maintenance of sexual size dimorphism among populations of three closely related lizard species (Phrynosoma douglasi, P. ditmarsi, and P. hernandezi). With independent contrasts I estimate evolutionary correlations among female body size, male body size, and sexual size dimorphism (SSD) to determine whether males have become small, females have become large, or both sexes have diverged concurrently in body size during the evolutionary Xhistory of this group. Population differences in degree of SSD are inversely correlated with average male body size, but are not correlated with average female body size. Thus, variation in SSD among populations has occurred predominantly through changes in male size, suggesting that selective pressures on small males may affect degree of SSD in this group. I explore three possible evolutionary mechanisms by which the mean male body size in a population could evolve: changes in size at maturity, changes in the variance of male body sizes, and changes in skewness of male body size distributions. Comparative analyses indicate that population differentiation in male body size is achieved by changes in male size at maturity, without changes in the variance or skewness of male and female size distributions. This study demonstrates the potential of comparative methods at lower taxonomic levels (among populations and closely related species) for studying microevolutionary processes that underlie population differentiation.     

Female‐biased dimorphism in size and age at maturity is reduced at higher latitudes in lake whitefish Coregonus clupeaformis

Female‐biased sexual dimorphism in size at maturity is a common pattern observed in freshwater fishes with indeterminate growth, yet can vary in magnitude among populations for reasons that are not well understood. According to sex‐specific optimization models, female‐biased sexual size dimorphism can evolve due to sexual selection favouring earlier maturation by males, even when sexes are otherwise similar in their growth and mortality regimes. The magnitude of sexual size dimorphism is expected to depend on mortality rate. When mortality rates are low, both males and females are expected to mature at older ages and larger sizes, with size determined by the von Bertalanffy growth equation. The difference between size at maturity in males and females becomes reduced when maturing at older ages, closer to asymptotic size. This phenomenon is called von Bertalanffy buffering. The predicted relationship between the magnitude of female‐biased sexual dimorphism in age and size at maturity and mortality rate was tested in a comparative analysis of lake whitefish Coregonus clupeaformis from 26 populations across a broad latitudinal range in North America. Most C. clupeaformis populations displayed female‐biased sexual dimorphism in size and age at 50% maturity. As predicted, female‐biased sexual size dimorphism was less extreme among lower mortality, high‐latitude populations.     

中国石龙子个体发育过程中头部两性异型和食性的变化

许多动物呈现个体大小、局部形态特征 (头部大小 )和体色的两性异形[5,14 ,15,2 1,2 2 ] 。 Darwin[12 ] 认为两性谋求各自最大的繁殖利益导致了两性异形 ,因此两性异形是性选择压力作用的结果。自 Darwin以来 ,许多同行认为性选择压力和非性选择压力均能导致动物的两性异形 ,两种选择压力在不同的动物中所起的作用是不同的 [2～ 5,7,10 ,16,2 1～ 2 6] 。性选择压力导致的两性异形与繁殖成功率直接有关。非性选择压力导致的两性异形与繁殖成功率无关或无直接的关系 ,如两性寿命的差异 [13 ]、两性食性的分离 [6,2 1]和两性分配用于生长的…     

A Genetic Test of Sexual Size Dimorphism in Pre-Emergent Chinook Salmon

Sex differences in early development may play an important role in the expression of sexual size dimorphism at the adult stage. To test whether sexual size dimorphism is present in pre-emergent chinook salmon (Oncorhynchus tshawytscha), alevins were reared at two temperatures (10 °C and 15 °C) and sexed using the OtY1 marker on the Y-chromosome. Linear mixed models were used to test for sex differences in alevin size within families while controlling for the random effects of sire and dam nested within sire. Males and females did not differ in weight at 10 °C but males were heavier than females at 15 °C. Sex accounted for 2% of the within-family variance in weight. In addition, at 15°C, the relationship between weight and sex was greater in families with larger eggs. Whereas male-biased sexual size dimorphism was present at the juvenile stage, female-biased sexual size dimorphism was present at sexual maturity. Males were also younger than females at sexual maturity. A head start on growth by males may underlie their earlier maturation at a smaller size, thus leading to female-biased SSD at the adult stage.     

Growth, size and age at maturity of the agile frog (Rana dalmatina) in an Iberian Peninsula population

The mean age of a population of agile frogs (Rana dalmatina) from the Iberian Peninsula was estimated using mark and recapture and skeletochronology. Life-history parameters, including growth rate, body length, age and size at maturity, sexual dimorphism and longevity, were studied. The regression between age and snout-vent length (SVL) was highly significant in both sexes. Males reached sexual maturity at two years of age, although sometimes they can reach it at only one year of age. The average SVL at maturity was 51.75 mm (standard error (SE) = 0.71; n = 45). Females reached sexual maturity at two years of age with an average SVL of 62.14 mm (SE = 2.20; n = 14). A subset of the female population reached sexual maturity at three years of age. Growth was rapid until sexual maturity was reached. There was an overlap of SVL between different age classes. Growth was continuous, fulfilling the conditions of Von Bertalanffy's model. The growth coefficient (K) was 0.840 in males and 0.625 in females. The maximum SVL was greater in females (73.00 mm) than in males (59.50 mm). Sexual dimorphism was significantly biased towards females in all age classes. The maximum longevity observed was 6 years in females and 8 years in males. Management strategies for agile frogs should take into account factors such as these life-history characteristics.     

Ontogeny of sexual dimorphism in the Long-tailed Manakin Chiroxiphia linearis: long maturation of display trait morphology

Males of dimorphic species often show ornaments that are thought to have evolved through female choice or/and male–male competition. The sexual differentiation of similar morphologies occurs during ontogeny, resulting in differential sex and age-specific selection. The Long-tailed Manakin is a dimorphic species with a highly skewed mating system, the males of which delay plumage maturation over 3 to 4 years. We describe ontogenetic changes in feather morphology in this species through sexual maturity. Males showed a significant increase in length of the central rectrices with age, hence their degree of sexual dimorphism increased from zero in 1-year-old males to 189.5% in adults. In contrast, male tail length decreased with age. Wing length did not vary significantly with age, but females had relatively longer wings than males. Wing loading was greater in females and decreased with age in males. In adults, rectrix length was positively correlated with testis volume, supporting the hypothesis that secondary sexual characters can signal the condition of primary sexual characters. Rectrix length showed positive allometry with body size in males less than 4 years old, whereas older males showed negative allometry and females showed isometry. Wing area and wing loading shifted from negative to positive allometry in males of 2 to 3 years of age. Changes in male morphology during ontogeny in the Long-tailed Manakin appeared to be associated with their specific display behaviours. Age-related changes in allometric growth of rectrices in the Long-tailed Manakin suggested that young males invest disproportionately more in the length of this trait relative to their body size. This investment could act as a signal of competitive ability to move status position in their orderly queue.     

Sexual size dimorphism,growth, and maturity of the fluvial eight-barbel loach in the Kako River,Japan

The maturation and growth pattern of the fluvial eight-barbel loach Lefua sp. (Japanese name: nagare-hotoke-dojo), an endangered species, was investigated using an individual identification-recapture method from 1995 to 1998 in an upper reach of a headwater tributary of the Kako River, Hyogo Prefecture, Japan. Based on observations of the gonads through the abdominal skin, the loach was estimated to breed mostly from May to July. All the males matured by age 1⁺, and all the females matured by age 2⁺. Gamete release in all individuals of both males and females was predicted from recaptured loaches during each breeding season. The standard length of mature females was significantly larger than that of males, showing sexual size dimorphism (SSD). The maximum sizes recorded were 75.4 mm SL for females and 61.2 mm SL for males. Both males and females of immature specimens grew mainly from May to November, including the breeding season, with no significant differences in growth rates between them. After sexual maturity, both males and females grew mainly from July to October (or November), after the breeding season, and the females exhibited higher growth rates than males. Therefore, SSD of the species seems to be attributable to the different growth rates after maturity. The longevity of the loach was estimated to exceed ten years based on individual growth patterns of various sizes during the survey period. It is likely that the loach has an iteroparous life history, breeding every year, and moderate growth rates after maturity.     

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.