Female-biased Sexual Size Dimorphism Research Articles

Parasite diversity of disease-bearing rodents of Southeast Asia: habitat determinants and effects on sexual size dimorphism and life-traits

We investigated a causal chain of relationships between habitat specialization and parasite species richness in rodent communities in Southeast Asia, and the consequences for variation in immune investment (using spleen size), the degree of sexual competition (using testes) and sexual size dimorphism (SSD). We used data gathered on rodents, their habitat specialization and their parasites (macro- and micro-parasites) in Southeast Asian landscapes. The results supported the hypotheses that parasite diversity drives the evolution of host life-traits and sexual selection. Firstly host habitat specialization explained the variation in parasite species richness. Secondly high parasite species richness was linked to host immune investment, using the relative spleen size of rodents. Thirdly according to the potential costs associated with immune investment, the relative spleen size was found to be negatively correlated with the relative size of testes among rodents. Fourthly, a positive relationship between male-biased SSD and parasite species richness was observed supporting the role of parasitism in sexual selection. Finally, the variation in SSD was positively associated with the degree of habitat specialization. Highest values of female-biased SSD were associated with habitat specialization, whereas highest values of male-biased SSD concerned synanthropic or generalist rodent species. These results, also correlative, will help to facilitate selection of the species that should be thoroughly investigated at the population level to better understand the selective effects of parasites on rodent life-history and behavior.

Sex-biased parental care and sexual size dimorphism in a provisioning arthropod

The diverse selection pressures driving the evolution of sexual size dimorphism (SSD) have long been debated. Whilst the balance between fecundity selection and sexual selection has received much attention, explanations based on sex-specific ecology have proven harder to test. In ectotherms, females are typically larger than males, and this is frequently thought to be because size constrains female fecundity more than it constrains male mating success. However, SSD could additionally reflect maternal care strategies. Under this hypothesis, females are relatively larger where reproduction requires greater maximum maternal effort—for example where mothers transport heavy provisions to the nests. To test this hypothesis, we focussed on digger wasps (Hymenoptera: Ammophilini), a relatively homogeneous group in which only females provision offspring. In some species, a single large prey item, up to ten times the mother’s weight, must be carried to each burrow on foot; other species provide many small prey, each flown individually to the nest. We found more pronounced female-biased SSD in species where females carry single, heavy prey. More generally, SSD was negatively correlated with the numbers of prey provided per offspring. Females provisioning multiple small items had longer wings and thoraxes, probably because smaller prey are carried in flight. Despite much theorising, few empirical studies have tested how sex-biased parental care can affect SSD. Our study reveals that such costs can be associated with the evolution of dimorphism, and this should be investigated in other clades where parental care costs differ between sexes and species.

The evolution of fecundity is associated with female body size but not female-biased sexual size dimorphism among frogs.

Sexual size dimorphism (SSD) is one of the most common ways in which males and females differ. Male-biased SSD (when males are larger) is often attributed to sexual selection favouring large males. When females are larger (female-biased SSD), it is often argued that natural selection favouring increased fecundity (i.e. larger clutches or eggs) has coevolved with larger female body size. Using comparative phylogenetic and multispecies regression model selection approaches, we test the hypothesis that among-species variation in female fecundity is associated with the evolution of female-biased SSD. We also ask whether the hypothesized relationship between SSD and fecundity is relaxed upon the evolution of parental care. Our results suggest a strong relationship between the evolution of fecundity and body size, but we find no significant relationship between fecundity and SSD. Similarly, there does not appear to be a relationship between fecundity and the presence or absence of parental care among species. Thus, although female body size and fecundity coevolve, selection for increased fecundity as an explanation for female-biased SSD is inconsistent with our analyses. We caution that a relationship between female body size and fecundity is insufficient evidence for fecundity selection driving the evolution of female-biased SSD.

Interpopulation Comparison of Sex-Biased Mortality and Sexual Size Dimorphism in Sea-Run Masu Salmon, Oncorhynchus masou.

Evolutionary ecologists often expect that natural and sexual selection result in systematic co-occurrence patterns of sex-biased mortality and sexual size dimorphism (SSD) within animal species. However, whether such patterns actually occur in wild animals is poorly examined. The following expectation, the larger sex suffers higher mortality, was primarily tested here for apparently native sea-run masu salmon (Oncorhynchus masou) in three populations in Hokkaido, Japan. Field surveys on sex ratios, body sizes, and ages of smolts and returning adults revealed that two of the three populations exhibited an expected pattern, a female-biased marine mortality and SSD, but one population demonstrated an unexpected co-occurrence of male-biased marine mortality and female-biased SSD. These female-biased SSDs were attributed to faster marine growth of females because of no sex difference in smolt body size. It has been previously suggested that breeding selection favoring large size generally act more strongly in females than in males in Japanese anadromous masu, as there is a weak sexual selection on adult males but universally intensive natural selection on adult females. Thus, this hypothesis explains female-biased SSDs well in all study populations. Interpopulation variation in sex-biased mortality found here might result from differences in marine predation and/or fishing pressures, given that selection driving female-biased SSD makes females forage more aggressively than males during the marine phase. Taken together, these results raise the possibility that evolutionary forces have shaped adaptive sex-specific foraging strategies under relationships between growth and mortality, resulting in co-occurrence patterns of sex-biased mortality and SSD within animal species.

Phenotypic Variation and Sexual Size Dimorphism in Dichroplus elongatus (Orthoptera: Acrididae).

Patterns of body size evolution are of particular interest because body size can affect virtually all the physiological and life history traits of an organism. Sexual size dimorphism (SSD), a difference in body size between males and females, is a widespread phenomenon in insects. Much of the variation in SSD is genetically based and likely due to differential selection acting on males and females. The importance of environmental variables and evolutionary processes affecting phenotypeic variation in both sexes may be useful to gain insights into insect ecology and evolution. Dichroplus elongatus Giglio-Tos is a South American grasshopper widely distributed throughout Argentina, Uruguay, most of Chile, and southern Brazil. In this study, we analyzed 122 adult females of D. elongatus collected in eight natural populations from central-east Argentina. Females show large body size variation among the analyzed populations and this variation exhibits a strong relationship with fecundity. Our results have shown that larger females were more fecund than smaller ones. We found that ovariole number varied along a latitudinal gradient, with higher ovariole numbers in populations from warmer locations. A considerable female-biased SSD was detected. SSD for three analyzed morphometric traits scaled isometrically. However, SSD for thorax length displayed a considerable variation across the studied area, indicating a larger relative increase in female size than in male size in warmer environmental conditions.

Effect of genomic deficiencies on sexual size dimorphism through modification of developmental time in Drosophila melanogaster.

Sexual size dimorphism (SSD), a difference in body size between sexes, is common in many taxa. In insects, females are larger than males in >70% of all taxa in most orders. The fruit fly, Drosophila melanogaster is one prominent model organism to investigate SSD since its clear and representative female-biased SSD and its growth regulation are well studied. Elucidating the number and nature of genetic elements that can potentially influence SSD would be helpful in understanding the evolutionary potential of SSD. Here, we investigated the SSD pattern caused by artificially introduced genetic variation in D. melanogaster, and examined whether variation in SSD was mediated by the sex-specific modification of developmental time. To map the genomic regions that had effects on sexual wing size and/or developmental time differences (SDtD), we reanalyzed previously published genome-wide deficiency mapping data to evaluate the effects of 376 isogenic deficiencies covering a total of ~67% of the genomic regions of the second and third chromosomes of D. melanogaster. We found genetic variation in SSD and SDtD generated by genomic deficiencies, and a negative genetic correlation between size and development time. We also found SSD and SDtD allometries that are not qualitatively congruent, which however overall at best only partly help in explaining the patterns found. We identified several genomic deficiencies with the tendency to either exaggerate or suppress SSD, in agreement with quantitative genetic null expectations of many loci with small effects. These novel findings contribute to a better understanding of the evolutionary potential of sexual dimorphism.

Geographic variation of life‐history traits in the sand lizard,Lacerta agilis: testing Darwin's fecundity‐advantage hypothesis

The fecundity-advantage hypothesis (FAH) explains larger female size relative to male size as a correlated response to fecundity selection. We explored FAH by investigating geographic variation in female reproductive output and its relation to sexual size dimorphism (SSD) in Lacerta agilis, an oviparous lizard occupying a major part of temperate Eurasia. We analysed how sex-specific body size and SSD are associated with two putative indicators of fecundity selection intensity (clutch size and the slope of the clutch size-female size relationship) and with two climatic variables throughout the species range and across two widespread evolutionary lineages. Variation within the lineages provides no support for FAH. In contrast, the divergence between the lineages is in line with FAH: the lineage with consistently female-biased SSD (L. a. agilis) exhibits higher clutch size and steeper fecundity slope than the lineage with an inconsistent and variable SSD (L. a. exigua). L. a. agilis shows lower offspring size (egg mass, hatchling mass) and higher clutch mass relative to female mass than L. a. exigua, that is both possible ways to enhance offspring number are exerted. As the SSD difference is due to male size (smaller males in L. a. agilis), fecundity selection favouring larger females, together with viability selection for smaller size in both sexes, would explain the female-biased SSD and reproductive characteristics of L. a. agilis. The pattern of intraspecific life-history divergence in L.agilis is strikingly similar to that between oviparous and viviparous populations of a related species Zootoca vivipara. Evolutionary implications of this parallelism are discussed.

Sex-specific phenotypic plasticity in response to the trade-off between developmental time and body size supports the dimorphic niche hypothesis

Female-biased sexual size dimorphism (SSD) is widespread in many invertebrate taxa. One hypothesis for the evolution of SSD is the dimorphic niche hypothesis, which states that SSD evolved in response to the different sexual reproductive roles. While females benefit from a larger body size by producing more or larger eggs, males benefit from a faster development, which allows them to fertilize virgin females (protandry). To test this hypothesis, we studied the influence of temperature and intraspecific density on the development of Chorthippus montanus. We reared them in monosexual groups under different conditions and measured adult body size, wing length, nymphal mortality, and development time. The present study revealed an inverse temperature–size relationship: body size increased with increasing temperature in both sexes. Furthermore, we found intersexual differences in the phenotypic response to population density, supporting the dimorphic niches hypothesis. At a lower temperature, female development time increased and male body size decreased with increasing density. Because there was no food limitation, we conclude that interference competition hampered development. By contrast to expectations, mortality decreased with increasing density, suggesting that interference did not negatively affect survival. The present study shows that sex-specific niche optima may be a major trigger of sexual dimorphisms. © 2015 The Linnean Society of London, Biological Journal of the Linnean Society, 2015, 115, 48–57.

Andrew meets Rensch: sexual size dimorphism and the inverse of Rensch's rule in Andrew's toad (Bufo andrewsi).

Variation in sexual size dimorphism (SSD) is a widespread phenomenon and is commonly attributed to variation in sex-specific patterns of selection. According to Rensch's rule, SSD increases with increasing body size when males are the larger sex, and decreases when females are the larger sex. Using data from 17 populations of Andrew's toad (Bufo andrewsi), we tested whether the patterns of SSD conform to Rensch's rule. Using field experiments, we also evaluated the hypothesis that sexual selection favours large male body size and that fecundity selection favours large female body size. The results revealed that the degree of SSD increased with increasing mean size in females, consistent with the inverse of Rensch's rule. Although experiments revealed evidence for a large-male mating advantage, selection for large male size was weak at best, and hence unlikely to be an important source of variation in SSD. However, fecundity selection favouring large females was evident, and likely to explain the observed inverse of Rensch's rule. After correcting male and female body size for age differences, the patterns of SSD remained the same, suggesting that the intra- and interpopulational variation in SSD is not driven by sex differences in age structure. Hence, these findings suggest that the strong fecundity selection favouring large females drives the evolution of female-biased SSD in B. andrewsi, providing an explanation for the inverse of Rensch's rule. As such, the study provides an important addition to the small body of literature that uses an intraspecific approach to demonstrate the inverse of Rensch's rule.

Size-dependent selective mechanisms on males and females and the evolution of sexual size dimorphism in frogs.

Sexual size dimorphism (SSD) varies in animals from male biased to female biased. The evolution of SSD is potentially influenced by a number of factors, such as territoriality, fecundity, and temporal breeding patterns (explosive vs. prolonged). In general, frogs show female-biased SSD with broad variance among species. Using comparative methods, we examine how different selective forces affect male and female sizes, and we test hypotheses about size-dependent mechanisms shaping SSD in frogs. Male size was weakly associated with SSD in all size classes, and we found no significant association among SSD, male size, temporal breeding pattern, and male territoriality. In contrast, female size best explained SSD variation across all size classes but especially for small-bodied species. We found a stronger evolutionary association between female body size and fecundity, and this fecundity advantage was highest in explosively breeding species. Our data indicate that the fecundity advantage associated with female body size may not be linear, such that intermediate and large females benefit less with body size increases. Therefore, size-dependent selection in females associated with fecundity and breeding patterns is an important mechanism driving SSD evolution in frogs. Our study underscores the fact that lineage-specific ecology and behavior should be incorporated in comparative analyses of animal SSD.

Sexual dimorphism and age of Mediterranean salamanders

We analysed sexual size dimorphism (SSD) for two Mediterranean species of the “true” salamander clade possessing distinct life histories (Salamandra algira and Mertensiella caucasica) and equilibrated the morphometric approach to individual age by using skeletochronology. For species that have a short breeding season and live at high altitudes, such as Mediterranean amphibians, the fecundity advantage hypothesis predicts female-biased SSD to maximise reproductive success. Our results showed no SSD in either species; however, morphometric data indicated a male-biased dimorphism in limb (arm and leg) dimensions in both species when compared to body size. Limb dimorphisms are likely related to the particular mating system, which involves an amplexus during spermatophore transfer. Arm length appeared sexually dimorphic during ontogeny both in viviparous S. algira and oviparous M. caucasica. A review on SSD indicated monomorphy of body size as a common lineage-specific pattern among the “true” salamander clade, but also the common presence of other traits such as sexually dimorphic limb proportions.

Sexual size dimorphism is not associated with the evolution of parental care in frogs.

Sex differences in parental care are thought to arise from differential selection on the sexes. Sexual dimorphism, including sexual size dimorphism (SSD), is often used as a proxy for sexual selection on males. Some studies have found an association between male-biased SSD (i.e., males larger than females) and the loss of paternal care. While the relationship between sexual selection on males and parental care evolution has been studied extensively, the relationship between female-biased SSD (i.e., females larger than males) and the evolution of parental care has received very little attention. Thus, we have little knowledge of whether female-biased SSD coevolves with parental care. In species displaying female-biased SSD, we might expect dimorphism to be associated with the evolution of paternal care or perhaps the loss of maternal care. Here, drawing on data for 99 extant frog species, we use comparative methods to evaluate how parental care and female-biased SSD have evolved over time. Generally, we find no significant correlation between the evolution of parental care and female-biased SSD in frogs. This suggests that differential selection on body size between the sexes is unlikely to have driven the evolution of parental care in these clades and questions whether we should expect sexual dimorphism to exhibit a general relationship with the evolution of sex differences in parental care.

Macroevolutionary patterns of sexual size dimorphism in copepods.

Major theories compete to explain the macroevolutionary trends observed in sexual size dimorphism (SSD) in animals. Quantitative genetic theory suggests that the sex under historically stronger directional selection will exhibit greater interspecific variance in size, with covariation between allometric slopes (male to female size) and the strength of SSD across clades. Rensch's rule (RR) also suggests a correlation, but one in which males are always the more size variant sex. Examining free-living pelagic and parasitic Copepoda, we test these competing predictions. Females are commonly the larger sex in copepod species. Comparing clades that vary by four orders of magnitude in their degree of dimorphism, we show that isometry is widespread. As such we find no support for either RR or for covariation between allometry and SSD. Our results suggest that selection on both sexes has been equally important. We next test the prediction that variation in the degree of SSD is related to the adult sex ratio. As males become relatively less abundant, it has been hypothesized that this will lead to a reduction in both inter-male competition and male size. However, the lack of such a correlation across diverse free-living pelagic families of copepods provides no support for this hypothesis. By comparison, in sea lice of the family Caligidae, there is some qualitative support of the hypothesis, males may suffer elevated mortality when they leave the host and rove for sedentary females, and their female-biased SSD is greater than in many free-living families. However, other parasitic copepods which do not appear to have obvious differences in sex-based mate searching risks also show similar or even more extreme SSD, therefore suggesting other factors can drive the observed extremes.

Phylogeny suggests nondirectional and isometric evolution of sexual size dimorphism in argiopine spiders.

Sexual dimorphism describes substantial differences between male and female phenotypes. In spiders, sexual dimorphism research almost exclusively focuses on size, and recent studies have recovered steady evolutionary size increases in females, and independent evolutionary size changes in males. Their discordance is due to negative allometric size patterns caused by different selection pressures on male and female sizes (converse Rensch's rule). Here, we investigated macroevolutionary patterns of sexual size dimorphism (SSD) in Argiopinae, a global lineage of orb-weaving spiders with varying degrees of SSD. We devised a Bayesian and maximum-likelihood molecular species-level phylogeny, and then used it to reconstruct sex-specific size evolution, to examine general hypotheses and different models of size evolution, to test for sexual size coevolution, and to examine allometric patterns of SSD. Our results, revealing ancestral moderate sizes and SSD, failed to reject the Brownian motion model, which suggests a nondirectional size evolution. Contrary to predictions, male and female sizes were phylogenetically correlated, and SSD evolution was isometric. We interpret these results to question the classical explanations of female-biased SSD via fecundity, gravity, and differential mortality. In argiopines, SSD evolution may be driven by these or additional selection mechanisms, but perhaps at different phylogenetic scales.

The evolution and maintenance of sexual size dimorphism in horseshoe crabs: an evaluation of six functional hypotheses

Ultimate hypotheses to explain sexual size dimorphism (SSD) generally focus on differences in reproductive roles between males and females. For example, intrasexual competition for mates can favour larger males in some taxa, and fecundity selection can favour larger females in other taxa. However, SSD is not simply due to one selection pressure on males or females, but is the outcome of numerous conflicting pressures on both sexes. Despite this, few studies have tested multiple hypotheses to explain the factors underlying SSD. Horseshoe crabs, Limulus polyphemus, have female-biased SSD in all populations, and are an interesting species for the study of SSD because of their unusual evolutionary history and life-history strategy. We used the hypothetico-deductive method to evaluate multiple predictions from six hypotheses to understand the evolution and maintenance of SSD in horseshoe crabs. We examined these hypotheses using field studies and data from the literature. We found little support for four hypotheses. However, predictions for the fecundity advantage hypothesis were supported (e.g. larger females lay more eggs than smaller females). Our results predict that, for every centimetre increase in size, females will lay 1672 ± 285 more eggs per spawning bout. Thus, selection is likely to favour the future compounded gain in fecundity from one extra year of growth over the gain from one additional mating opportunity if females mature earlier. Predictions for the protandry hypothesis were also supported. The costs for males maturing at smaller sizes (i.e. a large-size advantage for competing males, and increased ejaculate volume) appear to be minimal. Therefore, males that mature and reproduce earlier will likely be selected for, given the demographic advantage of their offspring hatching and reproducing earlier. Our study illustrates the importance of investigating multiple hypotheses for both males and females in the study of SSD.

Evolution and maintenance of sexual size dimorphism: aligning phylogenetic and experimental evidence

Integrating the insights derived from both phylogenetic and experimental approaches offers a more complete understanding of evolutionary patterns and processes, yet it is rarely a feature of investigations of the evolutionary significance of trait variation. We combine these approaches to reinterpret the patterns and processes in the evolution of female biased sexual size dimorphism in Nephilidae, a spider lineage characterized by the most extreme sexual size dimorphism among terrestrial animals. We use a molecular phylogeny to reconstruct the size evolution for each sex and reveal a case of “sexually dimorphic gigantism”: both sexes steadily outgrow their ancestral sizes, but the female and male slopes differ, and hence sexual size dimorphism steadily increases. A review of the experimental evidence reveals a predominant net selection for large size in both sexes, consistent with the phylogenetic pattern for females but not for males. Thus, while sexual size dimorphism in spiders most likely originates and is maintained by fecundity selection on females, it is unclear what selection pressures prevent males from becoming as large as females. This integrated approach highlights the dangers of inferring evolutionary significance from experimental studies that isolate the effects of single selection pressures.

Body Size Variation of European Storm PetrelsHydrobates pelagicusin Relation to Environmental Variables

Body size differentiation may have developed in response to environmental gradients. A pattern of large individuals prevailing in colder areas is often observed and is explained by the heat conservation hypothesis (Bergmann's rule). To understand patterns driving body size variation in a pelagic seabird, the European Storm Petrel Hydrobates pelagicus, we examined the relationship between wing length, body mass and environmental variables in breeding areas (sea surface temperature, air temperature and wind speed). As this species has been divided into two subspecies: Mediterranean H. p. melitensis and Atlantic H. p. pelagicus, we performed the analyses at different scales (species, Atlantic subspecies and regional North Atlantic). At the species and subspecies scales, there was a longitudinal increase in wing length from west to east. At the subspecies and regional scale, we found a latitudinal increase in this variable from south to north. This result and the significant increase of wing length with decreasing sea surface and air temperatures are concordant with Bergmann's rule. In addition, body mass at the species and subspecies scales decreased with increasing wind speed, what may have a functional implication (small body mass may increase manoeuvrability over waves in conditions of stronger wind). Both genetic (two subspecies differing in body size) and environmental factors seem to be important forces driving intercolony variation in body size. Our study on sexual size dimorphism (SSD) revealed that in 156 molecularly sexed adults from the Faeroes, wing and tail length, and body mass exhibited female-biased SSD, while head-bill length showed male-biased SSD. The best discriminant function for sexing based on body measurements correctly classified 75% of individuals. Considering low correctness of proposed functions and geographical variation of body size, use of alternative methods (e.g. molecular tools) is recommended for sex discrimination in the European Storm Petrel.

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.

Allometry for Sexual Size Dimorphism in Stoneflies Defies the Rensch's Rule.

The Rensch's rule predicts that male-biased sexual size dimorphism (SSD) increases with body size, while female-biased SSD (FBSSD) decreases. In insects, many groups follow this rule, but the evidence suggests that it is taxon dependent and that the inverse of the rule can occur in species with FBSSD. Therefore, we conducted this study with Gripopterygidae stoneflies (Plecoptera) to describe their pattern of SSD and determine if they follow the Rensch's rule. Our data suggest that stoneflies exhibit FBSSD and do not follow the rule, but a reverse pattern. Our results corroborate other studies that suggest that the Rensch's rule is taxon based and that groups with FBSSD usually fail to obey the rule.

A test of Rensch's rule in greater horseshoe bat (Rhinolophus ferrumequinum) with female-biased sexual size dimorphism.

Sexual size dimorphism (SSD) is widespread within the animal kingdom. Rensch’s rule describes a relationship between SSD and body size: SSD increases with body size when males are the larger sex, and decreases with body size when females are the larger sex. Rensch’s rule is well supported for taxa that exhibit male-biased SSD but patterns of allometry among taxa with female-biased size dimorphism are mixed, there is evidence both for and against the rule. Furthermore, most studies have investigated Rensch’s rule across a variety of taxa; but among-population studies supporting Rensch’s rule are lacking, especially in taxa that display only slight SSD. Here, we tested whether patterns of intraspecific variation in SSD in greater horseshoe bats conform to Rensch’s rule, and evaluated the contribution of latitude to Rensch’s rule. Our results showed SSD was consistently female-biased in greater horseshoe bats, although female body size was only slightly larger than male body size. The slope of major axis regression of log10 (male) on log10 (female) was significantly different from 1. Forearm length for both sexes of greater horseshoe bats was significantly negatively correlated with latitude, and males displayed a slightly but nonsignificant steeper latitudinal cline in body size than females. We suggest that variation in patterns of SSD among greater horseshoe bat populations is consistent with Rensch’s rule indicating that males were the more variable sex. Males did not have a steeper body size–latitude relationship than females suggesting that sex-specific latitudinal variation in body size may not be an important contributing factor to Rensch’s rule. Future research on greater horseshoe bats might best focus on more comprehensive mechanisms driving the pattern of female-biased SSD variation.