Sexual dimorphism in the human bony pelvis, with a consideration of the Neandertal pelvis from Kebara Cave, Israel.

Sexual dimorphism of the human pelvis is inferentially related to obstetrics. However, researchers disagree in the identification and obstetric significance of pelvic dimorphisms. This study addresses three issues. First, common patterns in dimorphism are identified by analysis of pelvimetrics from six independent samples (Whites and Blacks of known sex and four Amerindian samples of unknown sex). Second, an hypothesis is tested that the index of pelvic dimorphism (female mean x 100/male mean) is inversely related to pelvic variability. Third, the pelvic dimensions of the Neandertal male from Kebara cave, Israel are compared with those of the males in this study. The results show that the pelvic inlet is the plane of least dimorphism in humans. The reason that reports often differ in the identification of dimorphisms for this pelvic plane is that both the length of the pubis and the shape of the inlet are related to nutrition. The dimensions of the pelvis that are most dimorphic (that is, female larger than male) are the measures of posterior space, angulation of sacrum, biischial breadth, and subpubic angle. Interestingly, these dimensions are also the most variable. The hypothesis that variability and dimorphism are inversely related fails to be supported. The factors that influence pelvic variability are discussed. The Kebara 2 pelvis has a spacious inlet and a confined outlet relative to modern males, though the circumferences of both planes in the Neandertal are within the range of variation of modern males. The inference is that outlet circumference in Neandertal females is also small in size, but within the range of variation of modern females. Arguments that Neandertal newborns were larger in size than those of modern humans necessarily imply that birth was more difficult in Neandertals.     

Costal process of the first sacral vertebra: sexual dimorphism and obstetrical adaptation

The human sacrum is sexually dimorphic, with males being larger than females in most dimensions. Previous studies, though, suggest that females may have a longer costal process of the first sacral vertebra (S1) than males. However, these studies neither quantified nor tested statistically the costal process of S1. This study compares S1 with the five lumbar vertebrae (L1 to L5) for a number of metric dimensions, including costal process length. Four issues are addressed, the: 1) hypothesis that females have a longer costal process of S1 than males; 2)hypothesis that homologous structures (i.e., costal processes of L1 to S1) differ in their direction of sexual dimorphism; 3) importance of the costal process of S1 to the obstetrical capacity of the pelvis; and 4) evolution of sexual dimorphism in costal process length of S1. One hundred ninety-seven individuals, including males and females of American blacks and whites, from the Hamann-Todd and Terry Collections were studied. Results show that males are significantly larger than females for most vertebral measurements, except that females have a significantly longer costal process of S1 than males. Costal process length of S1 is positively correlated with the transverse diameter and circumference of the pelvic inlet. The magnitude of sexual dimorphism in costal process length of S1 ranks this measure among the most highly dimorphic of the pelvis. Compared with the humans in this study, australopithecines have a relatively long costal process of S1, but their broad sacrum was not associated with obstetrical imperatives.     

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.     

Pubic symphyseal synostosis and sexual dimorphism of the pelvis inPresbytis cristata andPresbytis rubicunda

This study examines the relationship between public symphyseal synostosis and sexual dimorphism of the pelvis in two sympatric species ofPresbytis—P. cristata andP. rubicunda. Whereas no specimen ofP. cristata shows fusion of the interpubic joint, a high percentage of female (43.8%) and male (83.3%)P. rubicunda have a fused public symphysis. As females of both species are similar in body size, they are predicted to give birth to similarly sized newborns. Based on comparison with other anthropoids, the percentage dimorphism in the ischiopubic index inP. cristata andP. rubicuda suggests selection on pelvic capacity in relation to obstetrics. In species characterized by cephalopelvic constriction (i.e., the size of the fetal cranium closely approximates the capacity of the maternal birth canal), successful birth seems possible only by a hormonally induced increase in pelvic joint mobility during delivery. However, fusion of the interpubic joint obviates pelvic joint mobility. Consequently, this study tests the hypothesis thatP. rubicunda shows obstetric adaptations of the pelvis that are not found inP. cristata. The results show that pelvic capacity is larger in females than males in bothP. cristata andP. rubicunda; the sexual difference is most pronounced at the inlet. Moreover, the pattern of pelvic dimorphism is nearly identical between the species. When females of the two species are compared,P. rubicunda evidences a shorter distance between the sacroiliac and hip joints and a wider bituberous diameter. The former is related to interspecific differences in locomotion, and the latter is associated with obstetrics.     

Adjustment of the fetal head and adult pelvis in modern humans

In nonhuman anthropoids, the anteroposterior (AP) diameters of the fetus are greater than the transverse (TR) diameters and the AP diameters of the pelvic planes are greater than the TR diameters: during labor, therefore, the fetus moves through the birth canal without changing position or orientation. In modern humans, the fetal head at term is encephalized and the fetal chest is flattened. The maternal pelvic inlet is flattened in an AP direction, the sacral promontory and the ischial spines are prominent. As a result, AP<TR at the inlet, but AP>TR at the midpelvis and outlet. In addition, the birth canal presents a marked sacral curvature in the AP direction. The human fetus successfully negotiates the birth canal because the three crucial fetal adaptations: (1) spheroidicity of the presenting part of the fetal head, which allows it to “roll” in the pelvis; (2) mobility of the head and chest in all directions; and (3) a capacity for cranial molding, which adapts fetal head dimensions to pelvic dimensions. The result is that the human fetal head and chest can perform multiple rotational movements in order to always present the greatest fetal diameters to the greatest pelvic diameters. Monkeys show a limited degree of encephalization and suffer from narrow TR pelvic diameters without any possibility of fetal adaptations as shown by humans. Apes also show some encephalization but, because of wider TR diameters in the pelvis, they achieve an easy delivery with no need of fetal adaptations.     

The obstetrical dilemma hypothesis: there's life in the old dog yet

The term ‘obstetrical dilemma’ was coined by Washburn in 1960 to describe the trade-off between selection for a larger birth canal, permitting successful passage of a big-brained human neonate, and the smaller pelvic dimensions required for bipedal locomotion. His suggested solution to these antagonistic pressures was to give birth prematurely, explaining the unusual degree of neurological and physical immaturity, or secondary altriciality, observed in human infants. This proposed trade-off has traditionally been offered as the predominant evolutionary explanation for why human childbirth is so challenging, and inherently risky, compared to that of other primates. This perceived difficulty is likely due to the tight fit of fetal to maternal pelvic dimensions along with the convoluted shape of the birth canal and a comparatively low degree of ligamentous flexibility. Although the ideas combined under the obstetrical dilemma hypothesis originated almost a century ago, they have received renewed attention and empirical scrutiny in the last decade, with some researchers advocating complete rejection of the hypothesis and its assumptions. However, the hypothesis is complex because it presently captures several, mutually non-exclusive ideas: (i) there is an evolutionary trade-off resulting from opposing selection pressures on the pelvis; (ii) selection favouring a narrow pelvis specifically derives from bipedalism; (iii) human neonates are secondarily altricial because they are born relatively immature to ensure that they fit through the maternal bony pelvis; (iv) as a corollary to the asymmetric selection pressure for a spacious birth canal in females, humans evolved pronounced sexual dimorphism of pelvic shape. Recently, the hypothesis has been challenged on both empirical and theoretical grounds. Here, we appraise the original ideas captured under the ‘obstetrical dilemma’ and their subsequent evolution. We also evaluate complementary and alternative explanations for a tight fetopelvic fit and obstructed labour, including ecological factors related to nutrition and thermoregulation, constraints imposed by the stability of the pelvic floor or by maternal and fetal metabolism, the energetics of bipedalism, and variability in pelvic shape. This reveals that human childbirth is affected by a complex combination of evolutionary, ecological, and biocultural factors, which variably constrain maternal pelvic form and fetal growth. Our review demonstrates that it is unwarranted to reject the obstetrical dilemma hypothesis entirely because several of its fundamental assumptions have not been successfully discounted despite claims to the contrary. As such, the obstetrical dilemma remains a tenable hypothesis that can be used productively to guide evolutionary research.     

Sexual dimorphism in the size and shape of the os coxae and the effects of microevolutionary processes

Sexual dimorphism in the human pelvis has been studied widely for forensic purposes, but it is still unclear to what extent it varies among human populations. There is evidence that microevolutionary processes, both neutral (i.e., population history) and selective (e.g., thermoregulatory adaptation and size‐related obstetrical constraints) contribute to explain pelvic variation among populations, but the extent to which these factors affect pelvic sexual dimorphism is unknown. In this study, I analyze sexual dimorphism of the os coxae in 20 globally distributed human populations, using 3D morphometric data to separate the size and shape components of sexual differences. After evaluating population differences in the degree and pattern of sexual dimorphism, I test for the effect of population history, climate, and body size in shaping global diversity. The results show that size and shape dimorphism follow different patterns. Coxal size dimorphism is generally quite consistent through populations, with males bigger than females, but it appears to be reduced in small‐bodied populations, possibly in relation to obstetrically‐related selective pressures for a spacious birth canal. Beyond a general species‐wide pattern of shape dimorphism, commonly used for forensic sex determination, other aspects of sexual differences in coxal shape vary among human populations, reflecting the effects of neutral demographic processes and climatic adaptation. Am J Phys Anthropol 153:167–177, 2014. © 2013 Wiley Periodicals, Inc.     

Big-bodied males help us recognize that females have big pelves

Schultz ([1949] Am. J. Phys. Anthropol. 7:401-424) presented a conundrum: among primates, sexual dimorphism of the pelvis is a developmental adjunct to dimorphism in other aspects of the body, albeit in the converse direction. Among species in which males are larger than females in body size, females are larger than males in some pelvic dimensions; species with little sexual dimorphism in nonpelvic size show little pelvic dimorphism. Obstetrical difficulty does not explain this relationship. The present study addresses this issue, evaluating the relationship between pelvic and femoral sexual dimorphism in 12 anthropoid species. The hypothesis is that species in which males are significantly larger than females in femoral size will have a higher incidence, magnitude, and variability of pelvic sexual dimorphism, with females having relatively larger pelves than males, compared with species monomorphic in femoral size. The results are consistent with the hypothesis. The proposed explanation is that the default pelvic anatomy in adulthood is that of the female; testosterone redirects growth from the default type to that of the male by differentially enhancing and repressing growth among the pelvic dimensions. Testosterone also influences sexual dimorphism of the femur. The magnitude of the pelvic response to testosterone is greater in species that are sexually dimorphic in the femur than in those that are monomorphic.     

Sex determination of prehistoric central California skeletal remains using discriminant analysis of the femur and humerus

A large sample (n = 370) of Central California prehistoric skeletal remains was analyzed for sexual dimorphism of long bones using nine femoral and nine humeral dimensions. Sex of all individuals was assessed using traits of the os pubis. Discriminant analysis was done separately for the robust Early Horizon sample and the Middle/Late Horizon sample. Use of multiple variables did not produce appreciably better results over the use of several of the best variables, analyzed singly. Attention is focused on measurements of maximum diameter femoral head, femoral bicondylar width, and diameter of the humeral head (transverse or vertical). These variables produce excellent separation of the sexes with about 90% accuracy for the Middle/Late Horizon sample. They have been overlooked in the recent literature in which the relative values of length versus midshaft dimensions are debated. The measurements found to be superior in this study are taken at the ends of the bones where durability of these regions is indicated by large sample sizes found in this study. We suggest workers redirect their focus from the midshaft to the ends of the long bones. The discriminant analyses presented here have greater accuracy than most claims for either long bone or cranial sex determination. These standards, based on two samples with differing robusticity, may have applicability for workers in other areas who lack large skeletal samples.     

通过骨盆外测量推算中骨盆径的一元和多元回归方程

本文以126例生育年龄非妊娠女工的髂结节间径、髂前上棘间径、大转子间径及坐骨结节间径为自变量估计因变量中骨盆横径,求出多元回归方程:=—6.89+0.05x_1+0.12x_2+0.23x_3+0.77x_4;以骶耻外径和骨盆倾斜度为自变量估计因变量中骨盆矢径,其回归方程为:=6.09+0.39x_1-0.05x_2。对两方程进行方差分析,其F值分别为4.069和8.26,均具高度显著性,可以应用于临床对中骨盆径的估计以取代X线测量法。     

Sexual dimorphism and mechanics of the human hip: A multivariate assessment

The present research was undertaken to determine the effects of sexual dimorphism in the human pelvis and femur on the mechanics of human locomotion. The analysis was based on six biomechanical variables determined from 25 male and 32 female skeletal remains from the Dickson Mound site. Discriminant function analysis indicates that the mechanical variables which primarily contribute to dimorphism are the moment arm of the gluteus medius and the torque produced by the abductors at the hip. These mechanical aspects of hip function produce greater pressure on the femoral head in females.     

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.

     

Bony pelvic canal size and shape in relation to body proportionality in humans

Obstetric selection acts on the female pelvic canal to accommodate the human neonate and contributes to pelvic sexual dimorphism. There is a complex relationship between selection for obstetric sufficiency and for overall body size in humans. The relationship between selective pressures may differ among populations of different body sizes and proportions, as pelvic canal dimensions vary among populations. Size and shape of the pelvic canal in relation to body size and shape were examined using nine skeletal samples (total female n = 57; male n = 84) from diverse geographical regions. Pelvic, vertebral, and lower limb bone measurements were collected. Principal component analyses demonstrate pelvic canal size and shape differences among the samples. Male multivariate variance in pelvic shape is greater than female variance for North and South Africans. High‐latitude samples have larger and broader bodies, and pelvic canals of larger size and, among females, relatively broader medio‐lateral dimensions relative to low‐latitude samples, which tend to display relatively expanded inlet antero‐posterior (A‐P) and posterior canal dimensions. Differences in canal shape exist among samples that are not associated with latitude or body size, suggesting independence of some canal shape characteristics from body size and shape. The South Africans are distinctive with very narrow bodies and small pelvic inlets relative to an elongated lower canal in A‐P and posterior lengths. Variation in pelvic canal geometry among populations is consistent with a high degree of evolvability in the human pelvis. Am J Phys Anthropol 151:88–101, 2013. © 2013 Wiley Periodicals, Inc.     

Sexual dimorphism,asymmetry, and the effect of reproduction on pelvis bone in the bank vole, <Emphasis Type="Italic">Myodes glareolus</Emphasis>

Sexual dimorphism in anatomical traits has been widely studied in animals. Although pelvis dimorphism was mostly studied in humans, it occurs also in many other mammalian species. Here, we investigated sexual dimorphism in the pelvis of the bank vole Myodes glareolus using individuals with known sex and reproductive status of females (parous vs nulliparous). The analyses revealed that the size and shape of pelvis differed significantly between sexes, as well as between nulliparous and parous females. In comparison with males, females had a significantly longer pelvis and pubis bones and a longer obturator foramen length, but a smaller pubis width. Interestingly, the difference between parous and nulliparous females resembles that between females and males: parous females had bigger pelvis, which probably resulted from changes during pregnancy and after birth. Left bones were on average larger than right ones, but the magnitude of directional asymmetry was not different between sex and reproduction group. Moreover, we noticed that fluctuating asymmetry of pelvis and pubis length was higher in females than in males and higher in parous than in nulliparous females, what is presumably associated with locomotor performance. A discriminant function analysis performed for the four bone size traits showed that the traits can be effectively used for a nearly perfect recognition of sexes and also a quite reliable recognition of the reproductive status of females.     

The secular trend in the diameter of the femur of American Whites and Negroes

The transverse diameter of the adult femur at the level of its mid-bicondylar length is related to length, age and year of birth in American White and Negro skeletons of both sexes of the Terry Collection. The coefficients of correlation between age at death and birth year are significant in all four sex-race groups, as are also the regression coefficients of transverse diameter on length and on age. The regression of length on age is negative and on birth year positive, although statistically significant only in the Negro male group. The partial regression coefficients of transverse diameter on length, on age, and on birth year, each eliminating the effects of the other two variables, are significant for length, not significant for age except in the Negro male group, and significant for birth year except in the Negro male group. Thus, the statistics of three of the sex-race groups do not support the hypothesis that transverse diameter of the femur increases with age, but rather suggest a negative secular trend; the apparent inconsistency in these findings of the Negro male group can, however, be considered insignificant from a statistical point of view.     

Sexual dimorphism in relative sacral breadth among catarrhine primates

As the sacrum contributes to the size and shape of the birth canal, the sexually dimorphic sacrum of humans is frequently interpreted within obstetric contexts. However, while the human sacrum has been extensively studied, comparatively little is known about sacral morphology in nonhuman primates. Thus, it remains unclear whether sacral sexual dimorphism exists in other primates, and whether potential dimorphism is primarily related to obstetrics or other factors such as body size dimorphism. In this study, sacra of Homo sapiens, Hylobates lar, Nasalis larvatus, Gorilla gorilla, Pongo pygmaeus, Pan troglodytes, and Pan paniscus were evaluated for sexual dimorphism in relative sacral breadth (i.e., the ratio of overall sacral breadth to first sacral vertebral body breadth). Homo sapiens, H. lar, N. larvatus, and G. gorilla exhibit dimorphism in this ratio. Of these, the first three species have large cephalopelvic proportions, whereas G. gorilla has small cephalopelvic proportions. P. pygmaeus, P. troglodytes, and P. paniscus, which all have small cephalopelvic proportions, were not dimorphic for relative sacral breadth. We argue that among species with large cephalopelvic proportions, wide sacral alae in females facilitate birth by increasing the pelvic inlet's transverse diameter. However, given the small cephalopelvic proportions among gorillas, an obstetric basis for dimorphism in relative sacral breadth appears unlikely. This raises the possibility that sacral dimorphism in gorillas is attributable to selection for relatively narrow sacra in males rather than relatively broad sacra in females. Accordingly, these results have implications for interpreting pelvic dimorphism among fossil primates, including hominins. Am J Phys Anthropol 152:435–446, 2013. © 2013 Wiley Periodicals, Inc.     

Sexual dimorphism of cusp dimensions in human maxillary molars

Cusp dimensions of human maxillary molars were compared between males and females to determine whether the later-developed, distal cusps displayed greater sexual dimorphism than the earlier-developed, mesial cusps, and whether the later-forming second molar displayed greater sexual dimorphism than the first molar. First and second permanent molar crowns (M1 and M2) were measured indirectly, using dental casts obtained from 117 Japanese (65 males and 52 females). Measurements included maximum mesiodistal and buccolingual crown diameters and the diameters of the four main cusps: the paracone, protocone, metacone, and hypocone. Mean values of crown dimensions were larger in males than in females for both M1 and M2, but the sexual difference in protocone diameter of M1 was not significant. The protocone in M1 showed the least amount of sexual dimorphism, followed by the metacone, hypocone, and paracone, while in M2, the percentage sexual dimorphism corresponded to the order of cusp formation: paracone, protocone, metacone, and hypocone. With the exception of the paracone diameter, M2 showed greater sexual dimorphism than M1. Sexual dimorphism was not always greater in the later-developed, distal cusps of M1 or M2, but the protocone, the most important cusp in terms of occlusal function, displayed the least dimorphism in M1.     

Tooth dimensions of Southern Chinese.

Teeth in casts of a random sample from the Hong Kong Oral Health Survey of 12-year-old children (n=459; 295 boys and 164 girls) were measured in the mesiodistal, buccolingual, and clinical crown height dimensions. Sexual dimorphism was evident in all tooth types in nearly all tooth dimensions, with the exception of the mesiodistal dimension of mandibular central incisors. The Chinese male tooth dimensions were larger than in females in nearly all characters. The measurements were compared with other human groups. Results showed that the Southern Chinese had larger tooth dimensions than the Japanese and than the White Americans. Hence it is important to have data concerning a relevant human group for purposes of clinical diagnosis and planning of treatment. These data may also be useful in forensic dentistry.     

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.     

When do meadow vipers (Vipera ursinii) become sexually dimorphic? – ontogenetic patterns of sexual size dimorphisms

Contrary to an increasing number of papers that document sexual dimorphism in size (and/or shape) in adults, studies dealing with sex differences in newborn and juvenile snakes are surprisingly scarce. Data about ontogenetic shifts in sexual dimorphism are generally lacking and hence, it is unclear whether sex differences are set at birth or arise post‐natally. In this study, we analyzed patterns of sexual dimorphism in body size, head dimensions and tail length (TL) among newborn, subadult and adult meadow vipers (Vipera ursinii) from the Bjelasica Mt. in Montenegro. Patterns of sexual size dimorphisms differed among traits. There was no significant difference in head dimension of males and females, but adult snakes were sexually dimorphic in body size. Sexual differences in TL were evident since birth but changed in degree throughout ontogeny. Neonate meadow vipers presented highly significant inter‐litter variation in the sexual dimorphism of all traits we have measured. Such family effects may have an important influence on extent of inter‐sexual differences in snakes and should be included in analyses of sexual dimorphism.