Partnership status and the temporal context of relationships influence human female preferences for sexual dimorphism in male face shape

Secondary sexual characteristics may indicate quality of the immune system and therefore a preference for masculinity may confer genetic benefits to offspring; however, high masculinity may be associated with costs of decreased paternal investment. The current study examined women's preferences for masculinity in male faces by using computer graphics to allow transformation between feminine and masculine versions of individual male faces. We found that preferences for masculinity are increased when women either have a partner or are considering a short-term relationship. Such preferences are potentially adaptive, serving to: (i) maximize parental investment and cooperation in long-term relationships by biasing choices towards feminine faced males, and (ii) maximize possible good-gene benefits of short-term or extra-pair partners by biasing choices towards masculine faced males. We also found that individuals using oral contraception do not show the above effects, indicating that such hormonal intervention potentially disrupts women's choices for evolutionarily relevant benefits from males.     

Sexual dimorphism in human skulls. A comparison of sexual dimorphism in different populations

Application and comparison of sex discriminant functions in different populations led to the conclusion that a certain combination and weighting of a few sex dimorphism variables (in this study we only used craniometric variables) can give a good discrimination between male and female individuals, independent of the racial group to which this function is applied. In our study, the sex-discriminatory power of five discriminant functions which were based on different ordination and selection procedures (e.g. professional knowledge, stepwise discriminant analysis, literature) of the cranial variables is compared. These discriminant functions were applied to three different data sets, the first being skull measurements from an Amsterdam series (Europids), the second skull measurements of a Zulu series (Negrids) and the third skull measurements of a Japan series (Mongolids). Our decision as to whether a function is a good or less good sex-discriminating function is determined by the Dt values (these values give an idea about the discriminatory value of the discriminant function when applied to a new test sample), the number of variables necessary to obtain this Dt and the location of the sectioning point (i.e. comparison between the estimation of the sectioning point and the ”real” sectioning point). These discriminant functions were compared withGiles Elliot's (1962, 1963) “race-independent” sex function.     

Statistical study of sexual dimorphism in the human fetal sciatic notch

Whether human fetal skeletal remains exhibit sexual dimorphism has been the subject of considerable debate. Most attention in this debate has focused on the greater sciatic notch of the ilium, since it is a gross morphological characteristic with known sex differences in the adult and is easily seen in fetal skeletal remains. Unfortunately, previous traditional morphometric analyses of the fetal sciatic notch have led to ambiguous results. The purpose of this study is to determine whether differences between the sexes can be discerned when modern morphometric techniques are applied to the fetal sciatic notch. Photographs of the ventral side of 133 fetal ilia of known age and sex from the Trotter Collection of Washington University were digitized, and the trace coordinates used for all subsequent analyses. The results of the analysis demonstrate that there is significant sexual dimorphism in the anterior to posterior location of the maximum depth of the sciatic notch, but that the depth of the notch itself is not dimorphic. While there is significant sexual dimorphism in the shape of the sciatic notch, the amount of overlap between males and females is too great for the sciatic notch to be used as a reliable indicator of sex. © 1995 Wiley-Liss, Inc.     

Mammalian sexual dimorphism

Sexual dimorphisms (SDs) have evolved in mammals to assure greater reproductive success for individuals, usually males. Secondary sexual characteristics (SSC) developed to further this objective, tending to be more pronounced in species which are polygynous, diurnal and open-habitat dwellers. Sexual selection has underpinned many of these changes, which are not necessarily advantageous for individual survival. Domestication has affected certain characteristics, more in terms of their quantitative rather than qualitative expression. However, restrictions imposed by domestication can also affect behaviors such as isolation and post-natal bonding while artificial selection can, by focusing on certain traits, cause unforeseen effects in genetically linked traits, which, when sex-specific or sex-linked, can be reflected in SD. On a global scale, environmental changes can have important phylogenetic implications for species which rely upon environmental cues for activities as migration, hibernation and breeding, especially when SD occurs in response to such cues. Understanding the evolutionary rationale behind the development of SDs, as well as the dynamics which occur at the interface between natural and artificial selection, allows positive insights into areas as diverse as wildlife preservation and livestock management. For both, greatest "success" should be achieved when artificial selection occurs in harmony with natural selection within a supportive environment. Thus the aim of this review is to discuss current knowledge relating to the evolution, benefits and costs of mammalian sexual dimorphisms and, where possible, draw conclusions that might be beneficial for the husbandry and propagation of mammals today.     

Visualizing sexual dimorphism in the brain

Sexually dimorphic behaviors are likely to involve neural pathways that express the androgen receptor (AR). We have genetically modified the AR locus to visualize dimorphisms in neuronal populations that express AR. Analysis of AR-positive neurons reveals both known dimorphisms in the preoptic area of the hypothalamus and the bed nucleus of the stria terminalis as well as novel dimorphic islands in the basal forebrain with a clarity unencumbered by the vast population of AR-negative neurons. This genetic approach allows the visualization of dimorphic subpopulations of AR-positive neurons along with their projections and may ultimately permit an association between neural circuits and specific dimorphic behaviors.     

Testing some hypotheses on human evolution and sexual dimorphism

Research on human evolution and sexual dimorphism motivates an interesting test problem. In studying hominid phylogeny it is of interest to test whether parallel evolution plays a role. With regard to sexual dimorphism it is of interest to known whether the directions of sexual dimorphism in the populations being compared are the same. We show that testing these two problems gives rise to the same type of hypothesis testing, viz. the problem of testing the hypothesis that the means of independent, normally distributed random vectors with unit covariance matrices are situated on a straight line through the origin. A test is proposed and applied to study the sexual dimorphism of 20 recent skull populations. In this example the hypothesis of equal directions of sexual dimorphism is rejected. The classical theory of constructing multiple discriminant functions (canonical variates) is adapted to the problem of comparing sexual dimorphisms.     

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.

     

A geometric morphometric study into the sexual dimorphism of the human scapula

Sex determination is vital when attempting to establish identity from skeletal remains. Two approaches to sex determination exists: morphological and metrical. The aim of this paper was to use geometric morphometrics to study the shape of the scapula and its sexual dimorphism.The sample comprised 45 adult black male and 45 adult black female scapulae of known sex. The scapulae were photographed and 21 homologous landmarks were plotted to use for geometric morphometric analysis with the ‘tps’ series of programs, as well as the IMP package. Consensus thin-plate splines and vector plots for males and females were compared. The CVA and TwoGroup analyses indicated that significant differences exist between males and females. The lateral and medial borders of females are straighter while the supraspinous fossa is more convexly curved than that of males. More than 91% of the females and 95% of the males were correctly assigned. Hotelling's T²-test yielded a significant p-value of 0.00039. In addition, 100 equidistant landmarks representing the curve only were also assigned. These, however, yielded considerably poorer results. It is concluded that it is better to use homologous landmarks rather than curve data only, as it is most probable that the shape of the outline relative to the fixed homologous points on the scapula is sexually dimorphic.     

Socio-bioenergetics and sexual dimorphism in primates

Socio-bioenergetics is presented as a practical method of estimating energy budgets of primates in a social context. Energy budgets are estimated on the basis of behavioral observations and a series of empirical formulae, which consider body weight, activity, and reproductive status. Data on a captive colony of Sykes' monkeys and baboons are incorporated as illustrations of the possible effects of group composition, body size, reproductive status, and activity patterns on energy requirements.Supported by the Wenner-Gren Foundation for Anthropological Research Incorporated and the National Science Foundation Grant GU-1598.     

On sexual dimorphism in immune function

Sexual dimorphism in immune function is a common pattern in vertebrates and also in a number of invertebrates. Most often, females are more 'immunocompetent' than males. The underlying causes are explained by either the role of immunosuppressive substances, such as testosterone, or by fundamental differences in male and female life histories. Here, we investigate some of the main predictions of the immunocompetence handicap hypothesis (ICHH) in a comparative framework using mammals. We focus specifically on the prediction that measures of sexual competition across species explain the observed patterns of variation in sex-specific immunocompetence within species. Our results are not consistent with the ICHH, but we do find that female mammals tend to have higher white blood cell counts (WBC), with some further associations between cell counts and longevity in females. We also document positive covariance between sexual dimorphism in immunity, as measured by a subset of WBC, and dimorphism in the duration of effective breeding. This is consistent with the application of 'Bateman's principle' to immunity, with females maximizing fitness by lengthening lifespan through greater investment in immune defences. Moreover, we present a meta-analysis of insect immunity, as the lack of testosterone in insects provides a means to investigate Bateman's principle for immunity independently of the ICHH. Here, we also find a systematic female bias in the expression of one of the two components of insect immune function that we investigated (phenoloxidase). From these analyses, we conclude that the mechanistic explanations of the ICHH lack empirical support. Instead, fitness-related differences between the sexes are potentially sufficient to explain many natural patterns in immunocompetence.     

On the origins of sexual dimorphism in butterflies

The processes governing the evolution of sexual dimorphism provided a foundation for sexual selection theory. Two alternative processes, originally proposed by Darwin and Wallace, differ primarily in the timing of events creating the dimorphism. In the process advocated by Darwin, a novel ornament arises in a single sex, with no temporal separation in the origin and sex-limitation of the novel trait. By contrast, Wallace proposed a process where novel ornaments appear simultaneously in both sexes, but are then converted into sex-limited expression by natural selection acting against showy coloration in one sex. Here, we investigate these alternative modes of sexual dimorphism evolution in a phylogenetic framework and demonstrate that both processes contribute to dimorphic wing patterns in the butterfly genera Bicyclus and Junonia. In some lineages, eyespots and bands arise in a single sex, whereas in other lineages they appear in both sexes but are then lost in one of the sexes. In addition, lineages displaying sexual dimorphism were more likely to become sexually monomorphic than they were to remain dimorphic. This derived monomorphism was either owing to a loss of the ornament ('drab monomorphism') or owing to a gain of the same ornament by the opposite sex ('mutual ornamentation'). Our results demonstrate the necessity of a plurality in theories explaining the evolution of sexual dimorphism within and across taxa. The origins and evolutionary fate of sexual dimorphism are probably influenced by underlying genetic architecture responsible for sex-limited expression and the degree of intralocus sexual conflict. Future comparative and developmental work on sexual dimorphism within and among taxa will provide a better understanding of the biases and constraints governing the evolution of animal sexual dimorphism.     

Development of sexual dimorphism in the embryonic gonad

Summary Differentiation of gonads in embryonic pigs (ages 24 to 44 days) was studied by light and electron microscopy. Incipient short cord-like structures were seen in embryos of both sexes at the age of 24 days. The cords were ultrastructurally similar. Medullary cords appeared in the testis at 26 days and in the ovary at 28 days. The cords in the ovary degenerated gradually and new cortical cords started to develop at 34 days. The formation of cords in both sexes is difficult to explain with the H-Y antigen theory, according to which the antigen should organize cords only in the male. The present results indicate that new approaches are needed for further development of the H-Y antigen theory.     

Statistics of sexual size dimorphism

In comparative studies of sexual size dimorphism (SSD), the methods used to quantify dimorphism are controversial. SSD is commonly expressed as a ratio between species mean values of males and females, such as M/F or (M-F)/([M+F]/2), but a number of investigators have suggested that ratios should not be used, mainly because their distributions usually violate the assumptions of parametric statistical tests, or because they lead to spurious relationships that invalidate the interpretation and statistical significance of regressions and correlations. As an alternative to ratios, the comparative study of SSD can be conducted by a combination of regression with sex-specific data and residuals from this regression. Twenty-five data sets were selected from the literature and used to duplicate a variety of statistical procedures commonly employed in studies of SSD. All analyses were repeated with five different ratios and with methods that avoid the calculation of any ratios. These data and a review of the statistical properties of ratios and residuals indicate that: (1) most of the ratios used in the SSD literature are unnecessary, and several commonly used ratios are statistically inferior to others. Only two ratios are needed, one on a logarithmic scale and one on a linear scale; (2) there is no problem with spurious correlation or non-normality when ratios are used in several types of statistical procedures commonly employed in studies of SSD; (3) residuals cannot replace ratios for the evaluation of many questions regarding the pattern of SSD among species; and (4) residuals usually are used incorrectly, leading to misspecified regression equations. Most of the questions for which residuals are used should be addressed by multiple regression. These results apply to studies using comparative methods with or without adjustments for phylogenetic effects.     

Ontogenetic approaches to sexual dimorphism in anthropoids

Studies of sexual dimorphism have traditionally focused on the static differences in size and shape between adult males and females. In this paper, I suggest that an investigation of the ontogenetic bases of sexual dimorphism can provide new insights and information unobtainable from studies concerned only with adult endpoints. While growth is often viewed as simply the developmental pathway utilized to attain final adult size and shape, we must recognize that it is the entire pattern of sex-differentiated growth, and not merely the adult endpoints, which is adaptive and the target of natural selection. The importance of an ontogenetic approach to the analysis of sexual dimorphism is also demonstrated by the fact that a given morphologicalresult (e.g., a certain degree of adult weight dimorphism) may be attained by very different developmentalprocesses, signalling selection for quite different factors. The need to analyze the ontogenetic bases of sexual dimorphism in size and shape has recently been recognized by Jarman, in his study of dimorphism in large terrestrial herbivores. Here I combine aspects of Jarman’s approach with those of allometry and heterochrony in an analysis of sexual dimorphism in selected anthropoid primates. It is demonstrated that although all dimorphic anthropoids appear to be characterized by somebimaturism, the degree varies significantly. Marked weight dimorphism in certain species is primarily produced by an increased differentiation of female and male growthrates, while in other species the primary change involves differences in thetime or duration of growth between the sexes. These variations are illustrated with anthropoid genera such asMiopithecus, Cercopithecus, Erythrocebus, Macaca, Papio, Pan, andGorilla. It is suggested that additional ontogenetic investigations of other anthropoids will help clarify some of the socioecological bases of this variation in the ways of attaining an adult dimorphic state. This will contribute to our understanding of the complex factors underlying and producing sexual dimorphism in primates and other mammals.     

The ontogeny of sexual dimorphism in the African apes

The relationship between the growth spurt and the onset of sexual maturity is problematic in nonhuman primates. Growth data on the cranium and postcranium of dentally aged pygmy chimpanzees, common chimpanzees, and gorillas are reported here. In all three species, male means generally exceed female means throughout growth, with the exception that females exhibit a spurt during one dental-age stage when they become generally larger than the males. This female spurt occurs earlier in an absolute and relative sense in the gorillas than the chimpanzees. These growth data support field and laboratory observations suggesting that female gorillas become sexually mature earlier than do female chimpanzees. Gorillas are thus characterized by a greater degree of “sexual bimaturism” than are the chimpanzees. Implications of these differences in terms of size dimorphism, mating systems, and morphology are discussed.