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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.

Biological Evolution↗

Sexual selection, natural selection and the evolution of dimorphic coloration and ornamentation in agamid lizards.

Both sexual selection and natural selection can influence the form of dimorphism in secondary sexual traits. Here, we used a comparative approach to examine the relative roles of sexual selection and natural selection in the evolution of sexually dimorphic coloration (dichromatism) and ornamentation in agamid lizards. Sexual dimorphism in head and body size were used as indirect indicators of sexual selection, and habitat type (openness) as an index of natural selection. We examined separately the dichromatism of body regions "exposed to" and "concealed from" visual predators, because these body regions are likely to be subject to different selection pressures. Dichromatism of "exposed" body regions was significantly associated with habitat type: males were typically more conspicuously coloured than females in closed habitats. By contrast, dichromatism of "concealed" body regions and ornament dimorphism were positively associated with sexual size dimorphism (SSD). When we examined male and female ornamentation separately, however, both were positively associated with habitat openness in addition to snout-vent length and head SSD. These results suggest that natural selection constrains the evolution of elaborate ornamentation in both sexes as well as sexual dichromatism of body regions exposed to visual predators. By contrast, dichromatism of "concealed" body regions and degree of ornament dimorphism appear to be driven to a greater degree by sexual selection.

Animals↗

Tissue-specific expression and regulation of sexually dimorphic genes in mice.

We report a comprehensive analysis of gene expression differences between sexes in multiple somatic tissues of 334 mice derived from an intercross between inbred mouse strains C57BL/6J and C3H/HeJ. The analysis of a large number of individuals provided the power to detect relatively small differences in expression between sexes, and the use of an intercross allowed analysis of the genetic control of sexually dimorphic gene expression. Microarray analysis of 23,574 transcripts revealed that the extent of sexual dimorphism in gene expression was much greater than previously recognized. Thus, thousands of genes showed sexual dimorphism in liver, adipose, and muscle, and hundreds of genes were sexually dimorphic in brain. These genes exhibited highly tissue-specific patterns of expression and were enriched for distinct pathways represented in the Gene Ontology database. They also showed evidence of chromosomal enrichment, not only on the sex chromosomes, but also on several autosomes. Genetic analyses provided evidence of the global regulation of subsets of the sexually dimorphic genes, as the transcript levels of a large number of these genes were controlled by several expression quantitative trait loci (eQTL) hotspots that exhibited tissue-specific control. Moreover, many tissue-specific transcription factor binding sites were found to be enriched in the sexually dimorphic genes.

Animals↗

Sexual size dimorphism in shorebirds, gulls, and alcids: the influence of sexual and natural selection.

Charadrii (shorebirds, gulls, and alcids) have an unusual diversity in their sexual size dimorphism, ranging from monomorphism to either male-biased or female-biased dimorphism. We use comparative analyses to investigate whether this variation relates to sexual selection through competition for mates or natural selection through different use of resources by males and females. As predicted by sexual selection theory, we found that in taxa with socially polygynous mating systems, males were relatively larger than females compared with less polygynous species. Furthermore, evolution toward socially polyandrous mating systems was correlated with decreases in relative male size. These patterns depend on the kinds of courtship displays performed by males. In taxa with acrobatic flight displays, males are relatively smaller than in taxa in which courtship involves simple flights or displays from the ground. This result remains significant when the relationship with mating system is controlled statistically, thereby explaining the enigma of why males are often smaller than females in socially monogamous species. We did not find evidence that evolutionary changes in sexual dimorphism relate to niche division on the breeding grounds. In particular, biparental species did not have greater dimorphism in bill lengths than uniparental species, contrary to the hypothesis that selection for ecological divergence on the breeding grounds has been important as a general explanation for patterns of bill dimorphism. Taken together, these results strongly suggest that sexual selection has had a major influence on sexual size dimorphism in Charadrii, whereas divergence in the use of feeding resources while breeding was not supported by our analyses.

Animals↗

Female-biased sexual size dimorphism in the yellow-pine chipmunk (Tamias amoenus): sex-specific patterns of annual reproductive success and survival.

Sexual size dimorphism is ultimately the result of independent, sex-specific selection on body size. In mammals, male-biased sexual size dimorphism is the predominant pattern, and it is usually attributed to the polygynous mating system prevalent in most mammals. This sole explanation is unsatisfying because selection acts on both sexes simultaneously, therefore any explanation of sexual size dimorphism should explain why one sex is relatively large and the other is small. Using mark-recapture techniques and DNA microsatellite loci to assign parentage, we examined sex-specific patterns of annual reproductive success and survival in the yellow-pine chipmunk (Tamias amoenus), a small mammal with female-biased sexual size dimorphism, to test the hypothesis that the dimorphism was related to sex differences in the relationship between body size and fitness. Chipmunks were monitored and body size components measured over three years in the Kananaskis Valley, Alberta, Canada. Male reproductive success was independent of body size perhaps due to trade-offs in body size associated with behavioral components of male mating success: dominance and running speed. Male survival was consistent with stabilizing selection for overall body size and body size components. The relationship between reproductive success and female body size fluctuated. In two of three years the relationship was positive, whereas in one year the relationship was negative. This may have been the result of differences in environmental conditions among years. Large females require more energy to maintain their soma than small females and may be unable to maintain lactation in the face of challenging environmental conditions. Female survival was positively related to body size, with little evidence for stabilizing selection. Sex differences in the relationship between body size and fitness (reproductive success and survival) were the result of different processes, but were ultimately consistent with female-biased sexual size dimorphism evident in this species.

Animals↗

Genetic constraints on floral evolution in a sexually dimorphic plant revealed by artificial selection.

Sexual dimorphism is one of the most widespread and recognizable patterns of phenotypic variation in the biotic world. Sexual dimorphism in floral display is striking in the dioecious plant Silene latifolia, with males making many, small flowers compared to females. We investigated this dimorphism via artificial selection on two populations to determine whether genetic variation exists within populations for flower size and the extent of the between-sex correlation, whether a flower size and number trade-off exists within each sex, and whether pollen and ovule production vary with flower size. We selected for decreased flower size (calyx width) in females and increased flower size in males and measured the response to selection in size and correlated responses in flower dry mass, flower number, and pollen or ovule number per flower. Four bouts of selection in each of two selection programs were performed, for a total of three selection lines to decrease size, three to increase it, and two control lines. Flower size always significantly responded to selection and we always found a significant correlated response in the sex not under selection. Selection decreased but did not eliminate the sexual dimorphism in flower dry mass and number. A negative relationship between flower size and number within each sex was revealed. Whereas ovule number showed a significant correlated response to selection on flower size, pollen number did not. Our results indicate that although substantial additive genetic variation for flower size exists, the high between-sex genetic correlation would likely constrain flower size from becoming more sexually dimorphic. Furthermore, floral display within each sex is constrained by a flower size and number trade-off. Given this trade-off and lack of variation in pollen production with flower size, we suggest that sexual dimorphism evolved via sexual selection to increase flower number in males but not females.

Biological Evolution↗

Sexual dimorphism in the postcranial skeleton of New World primates.

This study examines sexual dimorphism in 24 dimensions of the postcranial skeleton of four platyrrhine species: Callithrix jacchus, Saguinus nigricollis, Saimiri sciureus, and Cebus albifrons. The two callitrichid species show a relatively small amount of variation in the degree of sexual dimorphism among the different dimensions. Variation is considerably higher in the two cebid species as reflected by a mosaic pattern of sexual dimorphisms with males being significantly larger than females in some dimensions, and females significantly larger than males in others. In dimensions of the pectoral girdle and limb bones, males and females in each of the two cebid species are essentially scaled versions of each other, with males being peramorphic compared to females. This pattern is primarily the result of time hypermorphosis, i.e. an extension of the growth period in time in males. Rate hypermorphosis, i.e. an increase in the rate of growth in time in males, appears to play an additional role, however, in S. sciureus. By contrast, in dimensions of the true pelvis, sex differences in shape are dissociated from those in size. They are interpreted as the result of acceleration, i.e. increase in rate of shape change in females, as an adaptation to obstetrical functions. Interspecific analyses indicate positive allometry of mean degree of postcranial dimorphism with respect to body size. This coincides with previous findings by Leutenegger and Cheverud [1982, 1985] on the scaling of sexual dimorphism in body weight and canine size, and thus supports their model which posits selection on body size as the prime mover for the evolution of sexual dimorphism.

Aging↗

Alterations in sexually dimorphic biotransformation of testosterone in juvenile American alligators (alligator mississippiensis) from contaminated lakes.

The goal of this study was to determine whether hepatic biotransformation of testosterone is normally sexually dimorphic in juvenile alligators and whether living in a contaminated environment affects hepatic dimorphism. Lake Woodruff served as our reference site. Moonshine Bay, located on the west side of Lake Okeechobee, served as an intermediate site. Lake Apopka, the Belle Glade area located at the south end of Lake Okeechobee, and Water Conservation Area 3A, in the northern Everglades, served as our contaminated sites (all lakes are in Florida). Normal testosterone hydroxylase activity exhibited sexually dimorphic patterns of expression, with reference animals from Lake Woodruff exhibiting a female:male ratio of 1.44. This pattern was perturbed in all of the intermediate and contaminated sites investigated. Normal testosterone oxido-reductase activity exhibited sexually dimorphic expression (Lake Woodruff female:male ratio of 1.45). This pattern was altered in all contaminated sites investigated. UDP-Glucuronosyltransferase activity exhibited no sexually dimorphic pattern in animals collected from our reference site, with Lake Woodruff animals exhibiting a female:male ratio of 1.06. This pattern was perturbed in animals from Water Conservation Area 3A, which exhibited a female:male ratio of 0.65. Sulfotransferase activity demonstrated no sexual dimorphism at any of the sites investigated, although elevated activity was observed in males from the Lake Okeechobee watershed compared to those from Lake Woodruff. These data demonstrate different patterns of hepatic androgen biotransformation in animals living in contaminated environments.

Alligators and Crocodiles↗

Sexual dimorphism in body and brain weight and its association with paw preference in cats.

Sexual dimorphism in body and brain weight was studied in cats. Paw preference was assessed by food reaching test. Sexual dimorphism in body weight related to paw preference. Males weighted more than females in total sample and non-right-handed (NRH) cats (no significant difference in right-handed group). NRH males weighted more than right-handed (RH) males (no difference in females). Absolute total-brain weight did not show sexual dimorphism. Relative total-brain weight was sexually dimorphic only in NRH cats (females more than males). RH males weighted more than NRH males (no significant difference in females). Only absolute right-brain weighted more in males than females (total sample). Otherwise, no significant difference was found between absolute right- and left-brain weights of male and female cats. The relative right- and left-brain weighted more in females than males only in NRH cats (no sexual dimorphism in RH cats); more in RH males than NRH males; no difference in RH and NRH females. The overall results indicated that sexual dimorphism in body and brain weight is associated with cerebral lateralization in cats.

Animals↗

Establishing sexual dimorphism in humans.

Sexual dimorphism, i.e. the distinct recognition of only two sexes per species, is the phenotypic expression of a multi-stage procedure at chromosomal, gonadal, hormonal and behavioral level. Chromosomal--genetic sexual dimorphism refers to the presence of two identical (XX) or two different (XY) gonosomes in females and males, respectively. This is due to the distinct content of the X and Y-chromosomes in both genes and regulatory sequences, SRY being the key regulator Hormones (AMH, testosterone, Insl3) secreted by the foetal testis (gonadal sexual dimorphism), impede Müller duct development, masculinize Wolff duct derivatives and are involved in testicular descent (hormonal sexual dimorphism). Steroid hormone receptors detected in the nervous system, link androgens with behavioral sexual dimorphism. Furthermore, sex chromosome genes directly affect brain sexual dimorphism and this may precede gonadal differentiation.

Animals↗

[Sexual dimorphism of somatic characteristics of children of short stature].

The material of normal 199 Warsaw children with hypostatura previously elaborated from the point of view of morphological traits and biological maturity are analysed at present from the point of view of sexual dimorphism. Dimorphic values of the following traits were tested: body height, trunk length, lower extremity length, biacromial diameter, biiliac diameter, chest breadth, chest depth, distal femoral epiph., chest circumference, thigh circumference, arm circumference, subscapular skinfold and body weight. For a more detailed analysis sexual dimorphic index was evaluated. The following conclusions were drawn from this analysis: 1. Sex dimorphism in children with hypostatura is less distinctly expressed than in normal children. 2. The biggest dimorphic differences among analysed anthropologic traits in both age groups concerned the width of distal femoral epiph. 3. Sex dimorphic index of analysed somatic features is in parallel direction with the control group and is of higher value among 7 year old children with hypostatura. 4. Basing on the results, slower biological maturity was confirmed among children under study.

Age Factors↗

Sexual dimorphism and interspecific cranial form in two capuchin species: Cebus albifrons and C. apella.

Ontogenetic patterns of sexual dimorphism and cranial form in two capuchin monkeys, Cebus albifrons and C. apella, are investigated by means of univariate, bivariate, and multivariate statistics. The analyses are based on 23 linear variables. Univariate analyses indicate that similar ontogenetic patterns of cranial sexual dimorphism are present; however, interspecific differences exist in timing. Ontogenetic scaling is present in both species' crania; however, it is more prevalent in C. albifrons. Several departures are present in cranial regions associated with orbital shape, the dental arcade, and the muscles of mastication. The latter two indicate that sexual differences in diet and/or foraging strategies may exist. Sexual selection is suggested as being the primary selective regime underlying the observed patterns of cranial sexual dimorphism in each species. Interspecific comparisons confirm that C. apella possesses a more dimorphic cranium than C. albifrons and that sexual dimorphism in C. apella begins earlier in development. Although interspecific ontogenetic scaling is present in some cranial variables, C. apella is not just a scaled-up version of C. albifrons. These sympatric congeners seem to be differentiated by variables related to the orbital region and the masticatory apparatus, as indicated by both departures from ontogenetic scaling and results of the discriminant function analysis. Ecological selection, rather than varying degrees of sexual selection, is likely to be responsible for this finding given that C. apella is known to consume hard-object foods. This is consistent with the predicted outcome of the competitive exclusion principle.

Animals↗

Sexual dimorphism in the face of Australopithecus africanus.

Recently discovered crania of Australopithecus africanus from Sterkfontein Member 4 and Makapansgat enlarge the size range of the species and encourage a reappraisal of both the degree and pattern of sexual dimorphism. Resampling methodology (bootstrapping) is used here to establish that A. africanus has a greater craniofacial size range than chimpanzees or modern humans, a range which is best attributed to a moderately high degree of sexual dimorphism. Compared to other fossil hominins, this variation is similar to that of Homo habilis (sensu lato) but less than that of A. boisei. The finding of moderately high dimorphism is corroborated by a CV-based estimate and ratios between those specimens considered to be male and those considered to be female. Inferences about the pattern of craniofacial dimorphism in the A. africanus face currently rely on the relationship of morphology and size. Larger specimens, particularly Stw 505, show prominent superciliary eminences and glabellar regions, but in features related in part to canine size, such as the curvature of the infraorbital surface, large and small specimens of A. africanus are similar. In this respect, the pattern resembles that of modern humans more so than chimpanzees or lowland gorillas. A. africanus may also show novel patterns of sexual dimorphism when compared to extant hominines, such as in the form of the anterior pillar. However, males of the species do not exhibit characteristics of more derived hominins, such as A. robustus.

Animals↗

Univariate estimates of sexual dimorphism: the effects of intrasexual variability.

The difference between male and female values of quantitative traits depends on the distribution of the variables within each sex, increasing with the rise in the difference between male and female average values and with the decrease of the dispersion of measurements in both sexes. This paper deals with the sensitivity of some widely used indices (relative difference between male and female mean values (MDI), Student's t, and the so-called Bennett-Chakraborty-Majumder D coefficient) with respect to intrasexual variability. The Kolmogorov-Smirnov distance (KS) is suggested here as a further index of dimorphism, although it is not usually utilized for this purpose. The theoretical approach is accompanied by the analysis of empirical data (metric variables obtained from a sample of present Sardinians) and by computer simulations under various assumptions. Indices based on the difference between male and female average values are not able to evaluate fully the various aspects of dimorphism. Student's t proved to be an adequate measure of whole sex differences, both in real and in simulated samples, as intrasexual variability is included in its formulation. The D index also proved to be a good measure of undivided sexual dimorphism, as it is the result of formal examination, and from application to empirical or to simulated cases. The Kolmogorov-Smirnov distance gave the best performance both in formal examination and in the whole simulation results, as it takes into account intrasexual variability, and is applicable to any kind of distribution. In simulated cases it was sensitive to variations of means and variances, and it was able to evaluate variance dimorphism. Since the last three indices measure the combined effect of size and variance dimorphism, the joint use of the MDI index is suggested in order to isolate the relative contribution of the difference between the means.

Body Constitution↗

Patterns of joint size dimorphism in the elbow and knee of catarrhine primates.

Differences in body size between conspecific sexes may incur differences in the relative size and/or shape of load-bearing joints, potentially confounding our understanding of variation in the fossil record. More specifically, larger males may experience relatively greater limb joint stress levels than females, unless an increase in weight-related forces is compensated for by positive allometry of articular surface areas. This study examines variation in limb joint size dimorphism (JSD) among extant catarrhines to: 1) determine whether taxa exhibit JSD beyond that expected to simply maintain geometric similarity between sexes, and 2) test whether taxa differ in JSD (relative to body size dimorphism) with respect to differences in limb use and/or phylogeny. "Joint size" was quantified for the distal humerus and distal femur of 25 taxa. Analysis of variance was used to test for differences between sexes (in joint size ratios) and among taxa (in patterns of dimorphism). Multiple regression was used to examine differences in JSD among taxa after accounting for variation in body size dimorphism (BSD) and body size. Although degrees of humeral and femoral JSD tend to be the same within species, interspecific variation exists in the extent to which both joints are dimorphic relative to BSD. While most cercopithecoids exhibit relatively high degrees of JSD (i.e., positive allometry), nonhuman hominoids exhibit degrees of JSD closer to isometry. These results may reflect a fundamental distinction between cercopithecoids and hominoids in joint design. Overall, the results make more sense (from a mechanical standpoint) when adjustments to BSD are made to account for the larger effective female body mass associated with bearing offspring. In contrast to other hominoids, modern humans exhibit relatively high JSD in both the knee and elbow (despite lack of forelimb use in weight support). Estimates of BSD based on fossil limb bones will vary according to the extant analogue chosen for comparison.

Animals↗

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 herbs, 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.

Body Height↗

Sexual dimorphism in two subspecies of Ethiopian Baboons (Papio Hamadryas) and their hybrids.

Sexual dimorphism in mammals has been related to such variables as absolute body size, ecology, and various aspects of social behavior. Attempts to relate dimorphism to any of these variables have necessarily used small heterogeneous samples to represent the relevant species. We are concerned by the inevitable exclusion of any measure of variability in dimorphism and the consequent impossibility of assessing the significance of observed interspecific differences. In this paper we describe aspects of sexual dimorphism in anubis, hamadryas, and hybrid baboons from Ethiopia. Samples are large enough to permit a measure of intrapopulational variability. Hamadryas baboons are more dimorphic than anubis in epigamic features, but not in postcanine dentition, nor, contrary to previous reports, in body weight or canine tooth size. Hybrid males are more hamadryas-like and hybrid females more anubis-like, as would be predicted by the proposed mechanism for the establishment of the hybrid zone, namely the capture of anubis females by hamadryas males.

Animals↗

Sexual dimorphism in the baboon facial skeleton.

Baboons exhibit marked sexual dimorphism in many aspects of their morphology. Dimorphism is especially pronounced in the face. We use finite-element analysis to investigate the ontogeny of sexual dimorphism in a cross-sectional sample of baboon (Papio sp.) faces. This method provides detailed quantitative information about size and shape changes at anatomical landmarks in the face during growth. Allometric results suggest that sexual dimorphism in facial size and shape is produced by ontogenetic scaling: males and females share a common ontogenetic trajectory. Analyses of growth in time, which complement allometric analyses, show that female growth slows much earlier than male growth, accounting for the differences between sexes. Local size and local shape follow similar patterns of growth, but changes in these variables are slower in females. Local and global facial size are much more dimorphic than local and global facial shape.

Animals↗