Inferential assessment of the MI index of sexual dimorphism: A comparative study with some other sexual dimorphism measures

In order to provide inferential support to the MI measure of sexual dimorphism we proposed for populations distributed as mixture models with two normal components, an interval estimate is constructed. There do not appear to exist measures of sexual dimorphism that possess inferential properties other than some statistics used with this purpose. The use of these sample functions in such a context as well as the purported inferential support of some other sexual dimorphism indices are discussed. A biological case study illustrates the distinct inferential conclusions that can be obtained when the indices here discussed and the one we proposed are considered.     

An analysis of cercopithecoid odontometrics. II. Relations between dental dimorphism, body size dimorphism and diet.

Odontometric, dietary, and body weight data were collected for a sample of 29 cercopithecoid species. Each species was assigned to one of three diet classes (frugivore, folivore, and omnivore) , and indices were constructed to estimate the extent of sexual dimorphism in body weight, postcanine area and incisor width in each of the species. Analysis proceeded by means of the analysis of covariance with the dental dimorphism indices as the dependent variables. Body weight dimorphism was not significantly related to either measure of dental dimorphism across the sample, and an analysis by diet alone revealed that omnivores show significantly higher dental dimorphism than do either of the other two diet classes. The relationship between this result and theories of sexual subniche differentiation is discussed.     

Pelvic dimorphism in relation to body size and body size dimorphism in humans

Many mammalian species display sexual dimorphism in the pelvis, where females possess larger dimensions of the obstetric (pelvic) canal than males. This is contrary to the general pattern of body size dimorphism, where males are larger than females. Pelvic dimorphism is often attributed to selection relating to parturition, or as a developmental consequence of secondary sexual differentiation (different allometric growth trajectories of each sex). Among anthropoid primates, species with higher body size dimorphism have higher pelvic dimorphism (in converse directions), which is consistent with an explanation of differential growth trajectories for pelvic dimorphism. This study investigates whether the pattern holds intraspecifically in humans by asking: Do human populations with high body size dimorphism also display high pelvic dimorphism? Previous research demonstrated that in some small-bodied populations, relative pelvic canal size can be larger than in large-bodied populations, while others have suggested that larger-bodied human populations display greater body size dimorphism. Eleven human skeletal samples (total N: male = 229, female = 208) were utilized, representing a range of body sizes and geographical regions. Skeletal measurements of the pelvis and femur were collected and indices of sexual dimorphism for the pelvis and femur were calculated for each sample [ln(M/F)]. Linear regression was used to examine the relationships between indices of pelvic and femoral size dimorphism, and between pelvic dimorphism and female femoral size. Contrary to expectations, the results suggest that pelvic dimorphism in humans is generally not correlated with body size dimorphism or female body size. These results indicate that divergent patterns of dimorphism exist for the pelvis and body size in humans. Implications for the evaluation of the evolution of pelvic dimorphism and rotational childbirth in Homo are considered.     

Quantitative measure of sexual selection with respect to the operational sex ratio: a comparison of selection indices

Despite numerous indices proposed to predict the evolution of mating systems, a unified measure of sexual selection has remained elusive. Three previous studies have compared indices of sexual selection under laboratory conditions. Here, we use a genetic study to compare the most widely used measures of sexual selection in natural populations. We explored the mating and reproductive successes of male and female bank voles, Clethrionomys glareolus, across manipulated operational sex ratios (OSRs) by genotyping all adult and pup bank voles on 13 islands using six microsatellite loci. We used Bateman's principles (Is and I and Bateman gradients) and selection coefficients (s' and beta') to evaluate, for the first time, the genetic mating system of bank voles and compared these measures with alternative indices of sexual selection (index of monopolization and Morisita's index) across the OSRs. We found that all the sexual selection indices show significant positive intercorrelations for both males and females, suggesting that Bateman's principles are an accurate and a valid measure of the mating system. The Bateman gradient, in particular, provides information over and above that of other sexual selection indices. Male bank voles show a greater potential for sexual selection than females, and Bateman gradients indicate a polygynandrous mating system. Selection coefficients reveal strong selection gradients on male bank vole plasma testosterone level rather than body size.     

禄丰古猿的两性差别

本文首先讨论了粗壮池猿化石标本的性别判断问题,然后把粗壮池猿和禄丰西瓦古猿分别与有关的现生猿类的两性差别进行比较研究,得出这两个进化系统性别差异的时代变化的结论。这样的结论支持禄丰西瓦古猿的系统地位的论证。     

Genetic variability of sexual size dimorphism in a natural population of<Emphasis Type="Italic">Drosophila melanogaster</Emphasis>: An isofemale-line approach

Most animal species exhibit sexual size dimorphism (SSD). SSD is a trait difficult to quantify for genetical purposes since it must be simultaneously measured on two kinds of individuals, and it is generally expressed either as a difference or as a ratio between sexes. Here we ask two related questions: What is the best way to describe SSD, and is it possible to conveniently demonstrate its genetic variability in a natural population? We show that a simple experimental design, the isofemale-line technique (full-sib families), may provide an estimate of genetic variability, using the coefficient of intraclass correlation. We consider two SSD indices, the female-male difference and the female/male ratio. For two size-related traits, wing and thorax length, we found that both SSD indices were normally distributed. Within each family, the variability of SSD was estimated by considering individual values in one sex (the female) with respect to the mean value in the other sex (the male). In a homogeneous sample of 30 lines ofDrosophila melanogaster, both indices provided similar intraclass correlations, on average 0.21, significantly greater than zero but lower than those for the traits themselves: 0.50 and 0.36 for wing and thorax length respectively. Wing and thorax length were strongly positively correlated within each sex. SSD indices of wing and thorax length were also positively correlated, but to a lesser degree than for the traits themselves. For comparative evolutionary studies, the ratio between sexes seems a better index of SSD since it avoids scaling effects among populations or species, permits comparisons between different traits, and has an unambiguous biological significance. In the case ofD. melanogaster grown at 25?C, the average female/male ratios are very similar for the wing (1.16) and the thorax (1.15), and indicate that, on average, these size traits are 15–16% longer in females.     

Quantifying reciprocity in distylous and tristylous plant populations

Reciprocal herkogamy (heterostyly) is an example of extreme relevance of accuracy of the location of the sexual organs within some floral systems. It involves the reciprocal positioning of anthers and stigmas in flowers of different plants within the same population, and the accuracy of this positioning is important to promote out‐cross pollination, prevent self‐interference, or both. Hence, several indices have been proposed to quantify reciprocity, most of them for populations with two different morphs (distylous). Here, we propose an extension of our index of reciprocity for distylous populations to be applied also to populations with three morphs (tristylous), allowing effective comparisons of reciprocity between tristylous, but also distylous populations. As for the distylous version, the index is based on comparison of the position of every single sexual organ in the sample with each and every organ of the opposite sex, for each of the three possible organ levels. Due to the massive amount of calculations required, a macro was developed that is available as Supplementary Information and at the website of the authors. The index and macro were tested on several hypothetical tristylous and distylous populations with predetermined mean and dispersion of sexual organs at each level, as well as on several actual tristylous and distylous populations. The index proposed is a solid tool for the study of reciprocity in distylous and tristylous populations. Comparisons between distylous and tristylous populations are easily performed and can be readily interpreted. The applicability of the index is facilitated through the software provided.     

Sexual size dimorphism in anurans

Several hypotheses have been proposed to explain the direction and extent of sexual size dimorphism in anurans (in which males are usually smaller than females) as a result of sexual selection. Here, we present an analysis to test the hypothesis that sexual dimorphism in anurans is largely a function of differences between the sexes in life-history strategies. Morphological and demographic data for anurans were collected from the literature, and the mean size and age in each sex were calculated for 51 populations, across 30 species and eight genera. Comparisons across 14 Rana species, eight Bufo species and across the genera showed a highly significant relationship between size dimorphism, measured using the female-male size ratio, and mean female-male age difference. A comparison of a subset of 17 of these species for which phylogenetic information was available, using the method of independent contrasts, yielded a similar result. These results indicate that most of the variation in size dimorphism in the anura can be explained in terms of differences in the age structure between the sexes in breeding populations. If sexual selection has an effect on size dimorphism in anurans, it is likely to be only a secondary one.     

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. Am J Phys Anthropol 109:501–508, 1999. © 1999 Wiley-Liss, Inc.     

Sexual dimorphism in modern human permanent teeth

On average, males possess larger tooth crowns than females in contemporary human populations, although the degree of dimorphism varies within different populations. In previous studies, different amounts of either enamel or dentine were implicated as the cause of this dimorphism. In this study, we attempt to determine the nature of sexual dimorphism in the crowns of permanent modern human teeth and to determine if two contrasting tooth types (permanent third molars and canines) show identical patterns of dimorphism in enamel and dentine distribution. We estimated the relative contributions of both enamel and dentine to total crown size, from buccolingual sections of teeth. Our sample consisted of a total of 144 mandibular permanent third molars and 25 permanent mandibular canines of known sex. We show that sexual dimorphism is likely due, in part, to the presence of relatively more dentine in the crowns of male teeth. However, whatever the underlying cause, dimorphism in both tooth root and tooth crown size should produce measurable dimorphism in tooth weight, though this has not been previously explored. Therefore, we provide some preliminary data that indicate the usefulness of wet tooth weight as a measure of sexual dimorphism. Both male permanent third molars and canines are significantly heavier than those of females. The weight dimorphism reported here for both classes of teeth may prove a useful finding for future forensic studies. In particular, weights of canines may be more useful as a means of sexing modern human skeletal material than linear or area measurements of teeth.     

A new approach to the quantification of degree of reciprocity in distylous (sensu lato) plant populations

Background and Aims

Although evolution of sexual polymorphism has been traditionally analysed using discrete characters, most of these polymorphisms are continuous. This is the case of heterostyly. Heterostyly is a floral polymorphism successfully used as a model to study the evolution of the sexual systems in plants. It involves the reciprocal positioning of anthers and stigmas in flowers of different plants within the same population. Studies of the functioning of heterostyly require the quantification of the degree of reciprocity between morphs of heterostylous species. Some reciprocity indices have been proposed previously, but they show significant limitations that need to be dealt with. This paper analyses these existing indices, and proposes a new index that aims to avoid their main problems (e.g. takes into account population variability and offers a single value per population).

Methods

The new index is based on the comparison of the position of every single sexual organ in the population with each and every organ of the opposite sex. To carry out all the calculations, a macro was programmed with MS^®Visual Basic in MS^® Excel. The behaviour of the index is tested using hypothetical data to simulate different situations of dimorphic populations; the index is also tested with some actual populations of different species of the genus Lithodora.

Results and Conclusions

The index of reciprocity proposed here is a sound alternative to previous indices: it compares stigma–stamen height gaps for all potential crosses in the population, it comprises stigma–stamen distance as well as dispersion, it is not skewed by the more frequent sex, and it can be meaningfully compared between populations and species. It has produced solid results for both hypothetical and natural populations.Key words: Distyly, floral polymorphism, heterostyly, reciprocity index, stylar dimorphism, Lithodora     

No evidence for sexual dimorphism of facial width-to-height ratio in four large adult samples

Sexual dimorphism in physical appearance may be an important cue in both intra- and intersex competition. Recently, the facial width-to-height ratio (fWHR) has been proposed as a novel sexually dimorphic morphologic measure, with men suggested to have a higher fWHR than women. Currently, however, the status of fWHR as a sexually dimorphic trait is unclear. Here we tested for sexual dimorphism in fWHR, as well as in three additional, previously reported facial measures, in four (three Caucasian and one African) independent samples. In three of the four samples, no significant sex differences in fWHR were observed. In one sample, males showed a significantly lower (rather than higher) fWHR than females (this effect was no longer significant after controlling for body mass index). By contrast, significant and large sex differences were observed in all four samples for each of the three previously validated facial metrics, namely, (a) lower face/face height, (b) cheekbone prominence, and (c) face width/lower face height. These results provide strong evidence against the claim that fWHR, at least as measured from the surface of the face, is sexually dimorphic.     

Adult sex ratio,sexual dimorphism and sexual selection in a Mesozoic reptile

The evolutionary history of sexual selection in the geologic past is poorly documented based on quantification, largely because of difficulty in sexing fossil specimens. Even such essential ecological parameters as adult sex ratio (ASR) and sexual size dimorphism (SSD) are rarely quantified, despite their implications for sexual selection. To enable their estimation, we propose a method for unbiased sex identification based on sexual shape dimorphism, using size-independent principal components of phenotypic data. We applied the method to test sexual selection in Keichousaurus hui, a Middle Triassic (about 237 Ma) sauropterygian with an unusually large sample size for a fossil reptile. Keichousaurus hui exhibited SSD biased towards males, as in the majority of extant reptiles, to a minor degree (sexual dimorphism index −0.087). The ASR is about 60% females, suggesting higher mortality of males over females. Both values support sexual selection of males in this species. The method may be applied to other fossil species. We also used the Gompertz allometric equation to study the sexual shape dimorphism of K. hui and found that two sexes had largely homogeneous phenotypes at birth except in the humeral width, contrary to previous suggestions derived from the standard allometric equation.     

SEXUAL DIMORPHISM IN SPOTTED DOLPHINS (STENELLA ATTENUATA) IN THE EASTERN TROPICAL PACIFIC OCEAN

We analyzed 4 meristic and 32 morphometric cranial traits of 612 adult specimens of spotted dolphins ( Stenella attenuata ) from the eastern tropical Pacific Ocean for sexual dimorphism. Eleven 5" latitude/longitude blocks with five or more of each sex were assessed with a two-way ANOVA for sex and geographic differences. Interaction between these two factors was found for six measurements, suggesting that the degree of sexual dimorphism varies geographically for a few characters (although in no discernible geographic pattern). Sexual dimorphism was demonstrated for 23 of the 36 characters, with differences ranging from 0.00 to 5.88 percent. Females characteristically had a longer rostrum, while males generally had larger skulls overall. In terms of number of characters, the extent of sexual dimorphism demonstrated for skulls of spotted dolphins goes considerably beyond that shown for any other small delphine. A discriminant function involving a combination of 10 characters enabled us to identify correctly the sex of more than 75 percent of the specimens. A procedure for correcting specimen measurements is outlined that would enable an investigator to combine male and female specimens in geographic variation studies. A term (zwitter) is proposed for use when referring to specimens where measurements have been corrected to take into account differences between the sexes.     

Evolution of sexual size dimorphism in birds: test of hypotheses using blue tits in contrasted Mediterranean habitats

Many hypotheses, either sex‐related or environment‐related, have been proposed to explain sexual size dimorphism in birds. Two populations of blue tits provide an interesting case study for testing these hypotheses because they live in contrasting environments in continental France and in Corsica and exhibit different degree of sexual size dimorphism. Contrary to several predictions, the insular population is less dimorphic than the continental one but neither the sexual selection hypothesis nor the niche variation hypothesis explain the observed patterns. In the mainland population it is advantageous for both sexes to be large, and males are larger than females. In Corsica, however, reproductive success was greater for pairs in which the male was relatively small, i.e. pairs in which sexual size dimorphism is reduced. The most likely explanation is that interpopulation differences in sexual size dimorphism are determined not by sex‐related factors, but by differences in sex‐specific reproductive roles and responses to environmental factors. Because of environmental stress on the island as a result of food shortage and high parasite infestations, the share of parents in caring for young favours small size in males so that a reduced sexual size dimorphism is not the target of selection but a by‐product of mechanisms that operate at the level of individual sexes.     

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.     

Morphological variation in adult chimpanzees (Pan troglodytes verus) of the Taï National Park, Côte d'Ivoire

Twenty five adult chimpanzee skeletons (Pan troglodytes verus) of known age and sex (15 females, 10 males) from a long‐term study site in Taï National Park, Cote d'Ivoire present new data on variation. These skeletons provide a rare opportunity to measure the cranium and postcranium from the same individuals. We compare measurements and indices of the Taï sample with those of relatively complete Pan troglodytes schweinfurthii skeletons from Gombe National Park, Tanzania. Measurements of Pan paniscus are included as an outside comparison. The Taï and Gombe samples are analyzed by sex; combined sex samples are compared between the two groups, and the two sexes to each other. Taï females and males do not differ in most long bone lengths or in pelvic dimensions, but do differ significantly in cranial capacity, facial measurements, clavicle length, scapular breadth, and femur length. Gombe females and males differ significantly in some facial measurements and in scapular breadth. In combined sex samples, Taï individuals have lower cranial capacity, longer palate and mandible, and greater dimensions in the trunk and limb lengths. Taï females account for most of the variation; males differ from each other only in greater length of humerus and femur. The Taï skeletons provide new data for assessing individual variation and sexual dimorphism within and between populations and species. The combination of cranial and postcranial data provides a clearer picture of chimpanzee intraspecific and interspecific variation than can be gained from either data set alone. Am J Phys Anthropol, 2008. © 2007 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.

     

Quantitative analysis of sexual dimorphism and sex ratio in Hyphydrus ovatus (Coleoptera: Dytiscidae)

Sexual dimorphism in size, shape, and mass relative to length, and sex ratios are quantified for populations of Hyphydrus ovatus. a dytiscid beetle Males of H ovatus are not only longer than females, but also significantly larger in elytron length, thorax width, head width, leg length, total width, total depth, and abdominal segment length Two local populations differ slightly but significantly in total depth and abdominal segment length, but sexual dimorphism in size is similar for the two populations Hyphydrus ovatus are also sexually dimorphic in shape, with males having relatively broader heads and thoraxes than females The two populations differed slightly but significantly in relative abdominal segment length, but as with size, sexual dimorphism in shape is similar for the two populations Males are relatively heavier than females, although the slope of the log mass vs log length relationship is the same for the two sexes Sex ratios in field samples vary significantly over the summer, with percent females declining from c 50% to c 15% Sex ratios are significantly below 50% females m two of five monthly samples and in the total pooled set of samples Sexual dimorphism in size, shape, and relative mass, combined with male-biased sex ratios suggest that larger size of male H ovatus is a product of sexual selection