Sexual dimorphism in stature and women's work: a phylogenetic cross-cultural analysis.

The following cultural variables were tested for their association with sexual dimorphism: sexual division of labor, type of subsistence (hunting and agriculture), and polygyny. The transmission of these traits among populations was investigated. All the traits were found to be associated with phylogeny, indicating that they are inherited from mother to daughter populations. A cross-cultural comparative method was used which controls for the statistical effects of similarity due to common ancestry (Galton's problem). Cross-cultural variation in sexual dimorphism in stature is negatively associated with women's contribution to subsistence. Women are taller, relative to men, in societies where women contribute more to food production. This may be because female nutritional status is better in these societies. No relationship was found between sexual dimorphism and other aspects of subsistence or polygyny. These results are discussed in relation to other studies of sexual dimorphism in modern and archaeological populations, and in relation to cross-cultural variation in sex-biased parental investment.     

中国现代人群两性身高差异分布及其影响因素

本文对中国现代人群的两性身高差异分布状况及其影响因素进行了分析。选用152处中国现代人群（含69处汉族人群和83处少数民族人群）的男、女性身高数据,计算两性身高差异指数,并对比该指数在南、北方汉族和少数民族人群间的分布差异,同时分析纬度、气候、体格大小与城乡环境因素对两性身高差异程度的影响。结果表明,中国男性的平均身高比女性高出约7.16%(4.72%~9.26%）;南、北方汉族和少数民族之间的两性身高差异程度相似,北方汉族和南方汉族两性身高差异程度相似,但北方少数民族的两性身高差异明显大于南方少数民族。此外,两性身高差异程度与纬度、气温年较差和年均风速呈低度线性正相关,与年均气温、年均降水量和年均相对湿度呈低度线性负相关,而与体格大小和城乡环境并无显著关联。这提示遗传和自然环境因素在中国现代人群两性身高差异的区域化演变中更趋主导性,而社会环境因素的影响程度相对较低。     

Human size evolution: no evolutionary allometric relationship between male and female stature

In many animal groups, sexual size dimorphism tends to be more pronounced in species with large body size. Similarly, in a previous cross-cultural analysis, male and female stature in humans were shown to be positively allometrically related, indicating a similar relationship where populations with larger stature were more dimorphic. In this study, we re-examine the hypothesis of an allometric relationship between the sexes using phylogenetic methodology. First, however, we tested whether there exist phylogenetic signals in male and female stature. Data on mean stature from 124 human populations was gathered from the literature. A phylogenetic test showed that male and female stature were significantly associated with phylogeny. These results indicate that comparative methods that to some degree incorporate genetic relatedness between populations are crucial when analyzing human size evolution in a cross-cultural context. Further, neither non-phylogenetic nor phylogenetic analyses revealed any allometric relationship between male and female stature. Thus, we found no support for the idea that sexual dimorphism increases with increasing stature in humans.     

QUANTITATIVE GENETICS OF SEXUAL DIMORPHISM IN HUMAN BODY SIZE

A classical data set is used to predict the effect of selection on sexual dimorphism and on the population means of three characters—stature, span, and cubit—in humans. Given selection of equal intensity, the population means of stature and of cubit should respond more than 60 times as fast as dimorphism in these characters. The population mean of span should also respond far more rapidly than dimorphism, but no numerical estimate of the ratio of these rates was possible. These results imply that sexual dimorphism in these characters can evolve only very slowly. Consequently, hypotheses about the causes of sexual dimorphism cannot be tested by comparing the dimorphism of different human societies. It has been suggested that primate sexual dimorphism may be an allometric response to selection for larger body size. We show that such selection can indeed generate sexual dimorphism, but that this effect is too weak to account for the observed relationship between dimorphism and body size in primates.     

Human fertility variation, size-related obstetrical performance and the evolution of sexual stature dimorphism

In several animal species, change in sexual size dimorphism is a correlated response to selection on fecundity. In humans, different hypotheses have been proposed to explain the variation of sexual dimorphism in stature, but no consensus has yet emerged. In this paper, we evaluate from a theoretical and an empirical point of view the hypothesis that the extent of sexual dimorphism in human populations results from the interaction between fertility and size-related obstetric complications. We first developed an optimal evolutionary model based on extensive simulations and then we performed a comparative analysis for a total set of 38 countries worldwide. Our optimization modelling shows that size-related mortality factors do indeed have the potential to affect the extent of sexual stature dimorphism. Comparative analysis using generalized linear modelling supports the idea that maternal death caused by deliveries and complications of pregnancy (a variable known to be size related) could be a key determinant explaining variation in sexual stature dimorphism across populations. We discuss our results in relation to other hypotheses on the evolution of sexual stature dimorphism in humans.     

Height and sexual dimorphism of stature among human societies

In this study, which is concerned with the varying degrees of sexual dimorphism of stature between human societies, adult male and female height measurements and male-female height ratios – the measure of sexual dimorphism – from 216 societies are statistically compared with several variables: marriage practices, protein availability, the presence of milking herds, settlement size, and climate. Our results indicate that while greater mean male height is associated with polygynous marriage, marriage practices did not exert an influence on the degree of sexual dimorphism of stature. On the other hand, the results suggest that while sexual dimorphism in height has a strong genetic component, dietary factors can influence the degree of dimorphism.     

Latitude and intersocietal variation of human sexual dimorphism of stature

In a recent article Alexander et al.(1979) argue that intersocietal variation in human sexual dimorphism of stature can be explained by differing degrees of male- male competition for mates found in different types of societal marriage systems. In this paper we reexamine their data and suggest that their results may be more simply explained by reference to the latitude of the societies in their sample. Our investigation indicates that latitude has a significant influence on the degree of societal sexual dimorphism of stature. Peoples of the arctic region and the short peoples of the equatorial regions are less dimorphic in stature than midlatitude peoples. Thus, we conclude that the sociobiological hypothesis presented by Alexander et al.cannot alone explain the intersocietal variation in sexual dimorphism of stature and that environmental factors must be taken into account in the explanation of such variation.This research was supported by a Faculty Research Grant from North Texas State University.     

The evolution of sexual size dimorphism in the house finch. II. Population divergence in relation to local selection

Recent colonization of ecologically distinct areas in North America by the house finch (Carpodacus mexicanus) was accompanied by strong population divergence in sexual size dimorphism. Here we examined whether this divergence was produced by population differences in local selection pressures acting on each sex. In a long-term study of recently established populations in Alabama, Michigan, and Montana, we examined three selection episodes for each sex: selection for pairing success, overwinter survival, and within-season fecundity. Populations varied in intensity of these selection episodes, the contribution of each episode to the net selection, and in the targets of selection. Direction and intensity of selection strongly differed between sexes, and different selection episodes often favored opposite changes in morphological traits. In each population, current net selection for sexual dimorphism was highly concordant with observed sexual dimorphism--in each population, selection for dimorphism was the strongest on the most dimorphic traits. Strong directional selection on sexually dimorphic traits, and similar intensities of selection in both sexes, suggest that in each of the recently established populations, both males and females are far from their local fitness optimum, and that sexual dimorphism has arisen from adaptive responses in both sexes. Population differences in patterns of selection on dimorphism, combined with both low levels of ontogenetic integration in heritable sexually dimorphic traits and sexual dimorphism in growth patterns, may account for the close correspondence between dimorphism in selection and observed dimorphism in morphology across house finch populations.     

Ancient-to-modern secular changes in Korean stature

Statural growth in human populations is a sensitive indicator of socio-economic well-being, and improvements in socio-economic status are reflected in secular increases in adult height. In the present study, we investigated the statures of historical Korean societies to show how stature changed over time. Applying Fujii's equation, derived from modern Japanese, to the measurement of femora removed from 15th- to 19th-century Joseon tombs, the average heights of Korean adults during the Joseon dynasty were estimated to be 161.1 ± 5.6 cm and 148.9 ± 4.6 cm for males and females, respectively. Plotting statures for successive historical societies against time revealed that Korean heights remained relatively unchanged through to the end of the 19th century, a pattern that differs from that seen in many Western countries in which stature transiently decreases after the Middle Ages. In contrast, a sharp increase in Korean stature was observed at the beginning of the 20th century, similar to trends seen in other nations (although exact timing varies in different countries). There were no accompanying changes of stature sexual dimorphism. The data reported in this study reflect the unique historical experience of Korea; the relative isolation of Joseon society, the late onset of modernization (at the end of the 19th century), and the later occurrence of industrialization (during the 1960s).     

Genetic differences among populations in sexual dimorphism: evidence for selection on males in a dioecious plant

Genetic variation among populations in the degree of sexual dimorphism may be a consequence of selection on one or both sexes. We analysed genetic parameters from crosses involving three populations of the dioecious plant Silene latifolia, which exhibits sexual dimorphism in flower size, to determine whether population differentiation was a result of selection on one or both sexes. We took the novel approach of comparing the ratio of population differentiation of a quantitative trait (Q(ST) ) to that of neutral genetic markers (F(ST) ) for males vs. females. We attributed 72.6% of calyx width variation in males to differences among populations vs. only 6.9% in females. The Q(ST) /F(ST) ratio was 4.2 for males vs. 0.4 for females, suggesting that selection on males is responsible for differentiation among populations in calyx width and its degree of sexual dimorphism. This selection may be indirect via genetic correlations with other morphological and physiological traits.     

The evolution of sexual size dimorphism in the house finch. III. Developmental basis

Sexual size dimorphism of adults proximately results from a combination of sexually dimorphic growth patterns and selection on growing individuals. Yet, most studies of the evolution of dimorphism have focused on correlates of only adult morphologies. Here we examined the ontogeny of sexual size dimorphism in an isolated population of the house finch (Carpodacus mexicanus). Sexes differed in growth rates and growth duration; in most traits, females grew faster than males, but males grew for a longer period. Sexual dimorphism in bill traits (bill length, width, depth) and in body traits (wing, tarsus, and tail length; mass) developed during different periods of ontogeny. Growth of bill traits was most different between sexes during the juvenile period (after leaving the nest), whereas growth of body traits was most sexually dimorphic during the first few days after hatching. Postgrowth selection on juveniles strongly influenced sexual dimorphism in all traits; in some traits, this selection canceled or reversed dimorphism patterns produced by growth differences between sexes. The net result was that adult sexual dimorphism, to a large degree, was an outcome of selection for survival during juvenile stages. We suggest that previously documented fast and extensive divergence of house finch populations in sexual size dimorphism may be partially produced by distinct environmental conditions during growth in these populations.     

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.     

Sexual dimorphism and speciation on two ecological coins: patterns from nature and theoretical predictions

Adaptive divergence of phenotypes, such as sexual dimorphism or adaptive speciation, can result from disruptive selection via competition for limited resources. Theory indicates that speciation and sexual dimorphism can result from identical ecological conditions, but co-occurrence is unlikely because whichever evolves first should dissipate the disruptive selection necessary to drive evolution of the other. Here, we consider ecological conditions in which disruptive selection can act along multiple ecological axes. Speciation in lake populations of threespine sticklebacks (Gasterosteus aculeatus) has been attributed to disruptive selection due to competition for resources. Head shape in sticklebacks is thought to reflect adaptation to different resource acquisition strategies. We measure sexual dimorphism and species variation in head shape and body size in stickleback populations in two lakes in British Columbia, Canada. We find that sexual dimorphism in head shape is greater than interspecific differences. Using a numerical simulation model that contains two axes of ecological variation, we show that speciation and sexual dimorphism can readily co-occur when the effects of loci underlying sexually dimorphic traits are orthogonal to those underlying sexually selected traits.     

Parabolic variation in sexual selection intensity across the range of a cold‐water pipefish: implications for susceptibility to climate change

While an understanding of evolutionary processes in shifting environments is vital in the context of rapid ecological change, one of the most potent selective forces, sexual selection, remains curiously unexplored. Variation in sexual selection across a species range, especially across a gradient of temperature regimes, has the potential to provide a window into the possible impacts of climate change on the evolution of mating patterns. Here, we investigated some of the links between temperature and indicators of sexual selection, using a cold‐water pipefish as model. We found that populations differed with respect to body size, length of the breeding season, fecundity, and sexual dimorphism across a wide latitudinal gradient. We encountered two types of latitudinal patterns, either linear, when related to body size, or parabolic in shape when considering variables related to sexual selection intensity, such as sexual dimorphism and reproductive investment. Our results suggest that sexual selection intensity increases toward both edges of the distribution and that the large differences in temperature likely play a significant role. Shorter breeding seasons in the north and reduced periods for gamete production in the south certainly have the potential to alter mating systems, breeding synchrony, and mate monopolization rates. As latitude and water temperature are tightly coupled across the European coasts, the observed patterns in traits related to sexual selection can lead to predictions regarding how sexual selection should change in response to climate change. Based on data from extant populations, we can predict that as the worm pipefish moves northward, a wave of decreasing selection intensity will likely replace the strong sexual selection at the northern range margin. In contrast, the southern populations will be followed by heightened sexual selection, which may exacerbate the problem of local extinction at this retreating boundary.     

Is sexual dimorphism in the femur a "population specific phenomenon"?

The patterns of sexual dimorphism as well as the differences in amount between the populations were studied on a sample of 162 male and 159 female left femora, which were classified as Zulu, Sotho, Xosa and South Africans of European extraction. Multivariate analyses revealed that even adjacent African tribes exhibit a different pattern of sexual dimorphism, but there were similarities between Zulu and European femora. Furthermore, relative size differences, i.e. shape, discriminated more clearly between the sexes than did absolute size. Bicondylar width yielded a statistically significant higher degree of sexual dimorphism in Europeans when compared to African populations. This finding was interpreted in terms of the biomechanical demands on the femur under different living conditions. On the other hand, sexual dimorphism of femoral length did not differ among the populations. This was unexpected since femoral length correlates highly with stature, which was reported to show a lesser degree of sexual dimorphism in Africans than in Europeans. Detailed analyses of the results of the present study led to suggest that different living conditions may affect bones in complex ways of which linear growth is only one aspect.     

Genetic constraints and the evolution of display trait sexual dimorphism by natural and sexual selection

The evolution of sexual dimorphism involves an interaction between sex-specific selection and a breakdown of genetic constraints that arise because the two sexes share a genome. We examined genetic constraints and the effect of sex-specific selection on a suite of sexually dimorphic display traits in Drosophila serrata. Sexual dimorphism varied among nine natural populations covering a substantial portion of the species range. Quantitative genetic analyses showed that intersexual genetic correlations were high because of autosomal genetic variance but that the inclusion of X-linked effects reduced genetic correlations substantially, indicating that sex linkage may be an important mechanism by which intersexual genetic constraints are reduced in this species. We then explored the potential for both natural and sexual selection to influence these traits, using a 12-generation laboratory experiment in which we altered the opportunities for each process as flies adapted to a novel environment. Sexual dimorphism evolved, with natural selection reducing sexual dimorphism, whereas sexual selection tended to increase it overall. To this extent, our results are consistent with the hypothesis that sexual selection favors evolutionary divergence of the sexes. However, sex-specific responses to natural and sexual selection contrasted with the classic model because sexual selection affected females rather than males.     

Human skin-color sexual dimorphism: a test of the sexual selection hypothesis

Applied to skin color, the sexual selection hypothesis proposes that male preference for light-skinned females explains the presence of light skin in areas of low solar radiation. According to this proposal, in areas of high solar radiation, natural selection for dark skin overrides the universal preference of males for light females. But in areas in which natural selection ceases to act, sexual selection becomes more important, and causes human populations to become light-skinned, and females to be lighter than males. The sexual selection hypothesis proposes that human sexual dimorphism of skin color should be positively correlated with distance from the equator. We tested the prediction that sexual dimorphism should increase with increasing latitude, using adult-only data sets derived from measurements with standard reflectance spectrophotometric devices. Our analysis failed to support the prediction of a positive correlation between increasing distance from the equator and increased sexual dimorphism. We found no evidence in support of the sexual selection hypothesis.     

Among‐population variation and correlations in sexually dimorphic traits of Silene latifolia

Abstract The degree of sexual dimorphism in a trait may be determined directly by disruptive selection, as well as by correlations with other traits under selection. We grew seeds from nine populations of the dioecious plant Silene latifolia in a common‐garden experiment to determine whether phenotypic variation and correlations existed for floral, leaf and resource allocation traits, and whether this variation had a genetic component. We also determined the traits which were sexually dimorphic, the degree of dimorphism, and whether it varied among populations. Seven traits exhibited among‐population variation and sexual dimorphism. Variation in the degree of dimorphism occurred only for two traits, suggesting that dimorphism may be evolving more slowly than trait means. Males had more, smaller flowers, shorter leaves, and allocated less of their total biomass to stems and more to leaves than females. Flower production was the most sexually dimorphic trait and was correlated with all measured traits. Most traits exhibited significant correlations between the sexes. The pattern of correlations and the degree of sexual dimorphism among traits lead us to suggest that intrasexual selection for an exaggerated floral display in males has indirectly led to sexual dimorphism in a host of other traits.     

Sexual dimorphism in relation to big-game hunting and economy in modern human populations

Postcranial skeletal data from two recent Eskimo populations are used to test David Frayer's model of sexual dimorphism reduction in Europe between the Upper Paleolithic and Mesolithic. Frayer argued that a change from big-game hunting and adoption of new technology in the Mesolithic reduced selection for large body size in males and led to a reduction in skeletal sexual dimorphism. Though aspects of Frayer's work have been criticized in the literature, the association of big-game hunting and high sexual dimorphism is untested. This study employs univariate and multivariate analysis to test that association by examining sexual dimorphism of cranial and postcranial bones of two recent Alaskan Eskimo populations, one being big-game (whale and other large marine mammal) hunting people, and the second being salmon fishing, riverine people. While big-game hunting influences skeletal robusticity, it cannot be said to lead to greater sexual dimorphism generally. The two populations had different relative sexual dimorphism levels for different parts of the body. Notably, the big-game hunting (whaling) Eskimos had the lower multivariate dimorphism in the humerus, which could be expected to be the structure under greatest exertion by such hunting in males. While the exertions of the whale hunting economic activities led to high skeletal robusticity, as predicted by Frayer's model, this was true of the females as well as the males, resulting in low sexual dimorphism in some features. Females are half the sexual dimorphism equation, and they cannot be seen as constants in any model of economic behavior. © 1993 Wiley-Liss, Inc.     

Methodological considerations on the study of sexual dimorphism in past human populations

The degree of sexual dimorphism in human populations is influenced by stress, social role and by labour division. However, studies on ethnographic populations provided contradictory results. Unfortunately, most of these studies were based on stature only, which, as we could observe in a survey on pre-protohistoric circum-Medirerranean samples, is a poor indicator of functionally related dimorphism. A number of skeletal measurements were examined: skull, stature, transverse trunk diameters, long bones length, circumference and section, in order to assess their usefulness as indicators of functionally related dimorphism. The best indicators were represented by section and circumference of the long bones of the limbs, followed by cross-shoulder breadth (biclavicular length), stature and limb bone length, facial measurements, cranial measurements and sacral breadth. From the methodological point of view, it was found that:

1. It is better to calculate the index of dimorphism for each trait or set of traits within each sample. Then a weighted average of all the available samples is taken. The index derived from pooling a number of samples does not make biological sense. In pooled samples the distinction of between versus within sample differences is obscured;
2. It is better to combine an index which is based on the difference between averages and one which takes variability into account, because variability can also be an aspect of sexual dimorphism;
3. It is better to apply some allometric correction to the measurements used. For instance, the log transformation produces clearer trends of differential dimorphism among the various traits.