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 in the Pelvis of <Emphasis Type="Italic">Microcebus</Emphasis>

Pelvic sexual dimorphism occurs in many anthropoid species and is often attributed to obstetric selection on female pelvic morphology. Few studies of pelvic dimorphism have included strepsirrhine taxa, which typically have relatively smaller infants than those of anthropoids. Because smaller female primates give birth to relatively larger infants, it is possible that the pelves of Microcebus, the smallest extant primate genus, will show some evidence of selection on obstetric adequacy. A comparison of adult female and neonatal body masses indicates that individual neonatal Microcebus are relatively large compared to adult female body mass, even though members of the taxon frequently produce twins. I examined variation in the bony pelvis within a sample of Microcebus. I measured specimens from a single locality, which probably represent 1 population. I measured 8 pelvic and 3 femoral variables to investigate skeletal size and pelvic size and shape dimorphism. Females significantly exceed males in absolute values of sacral width, pelvic height, pubic length, and distances from the pubic symphysis to the ischial tuberosity and points on the sacrum. Measurements of the femur are not significantly greater in females, suggesting that the pelvic differences are not due to skeletal size dimorphism. Significant pelvic shape or ratio differences, calculated via the geometric mean of 5 variables as the denominator, included greater relative pubic length and sacral width in females. Hence selection for obstetric adequacy may occur in the extremely small-bodied Microcebus.     

Variation in pelvic size between males and females

Selection intensity and phenotypic variability are inversely related. It has been hypothesized that, owing to opposing selection pressures on pelvic morphology in females between efficiency in locomotion and obstetric adequacy, female pelvic morphology is less variable than that in males. The hypothesis was supported based on data derived from observational methods of sexing pelves (Meindl et al., 1985). The hypothesis was tested in the present study based on a metrical analysis of the true pelvis. The results show that there are no sexual differences in pelvic variability. Consequently, while males and females are equally variable in the dimensions of the true pelvis, the visual cues that osteologists use to sex pelves are more variable in males.     

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.     

Obstetric dimensions of the true pelvis in a medieval population from Sudanese Nubia.

Functional analysis of the true pelvis (defined as that portion lying below and including the pelvic brim) was undertaken on a sample of 36 females from the Medieval site of Kulubnarti in Sudanese Nubia. Standard obstetric measurements were taken and compared to four additional prehistoric skeletal samples and to modern American standards for the same obstetric dimensions. Relative to the other prehistoric populations, the Kulubnarti pelves are smaller in most dimensions and, when compared to modern American standards, from one-third to one-half would be diagnosed as contracted in one or more planes. Given the meager, fluctuating resources of these Medieval Nubians' harsh desert environment, pelvic size reduction is a likely result of body size reduction as one biological response to nutritional stress (Mittler and Van Gerven, 1989; Moore et al., 1986; Van Gerven et al., 1981). It is argued, however, that size reduction created a high potential for either maternal-neonatal morbidity and mortality due to fetopelvic disproportion or neonatal loss due to low birth weight. In either case, it is suggested that the Kulubnarti population paid a significant biological price for this aspect of size reduction.     

Sexual dimorphism in the pelves of African lorises

The present study is the first describing sexual dimorphism in the pelves of prosimian primates. Various measurements and indices indicate that there is no significant sexual dimorphism in the pelves of African lorises (Perodicticus potto and Arctocebus calabarensis). The lack of even a moderate degree of sexual dimorphism can be interpreted as the result of a lack of marked sexual differences in body size and of absence of selective pressure for expansion of the birth canal, the latter due to the small size of the fetus at term in relation to the dimensions of the female pelvic inlet.     

Body size estimation of small‐bodied humans: Applicability of current methods

Body size (stature and mass) estimates are integral to understanding the lifeways of past populations.Body size estimation of an archaeological skeletal sample can be problematic when the body size or proportions of the population are distinctive. One such population is that of the Holocene Later Stone Age (LSA) of southern Africa, in which small stature (mean femoral length = 407 mm, n = 52) and narrow pelves (mean bi‐iliac breadth = 210 mm, n = 50) produce a distinctive adult body size/shape, making it difficult to identify appropriate body size estimation methods. Material culture, morphology, and culture history link the Later Stone Age people with the descendant population collectively known as the Khoe‐San. Stature estimates based on skeletal “anatomical” linear measures (the Fully method) and on long bone length are compared, along with body mass estimates derived from “morphometric” (bi‐iliac breath/stature) and “biomechanical” (femoral head diameter) methods, in a LSA adult skeletal sample (n = 52) from the from coastal and near‐coastal regions of South Africa. Indices of sexual dimorphism (ISD) for each method are compared with data from living populations. Fully anatomical stature is most congruent with Olivier's femur + tibia method, although both produce low ISD. McHenry's femoral head body mass formula produces estimates most consistent with the bi‐iliac breadth/staturemethod for the females, although the males display higher degrees of disagreement among methods. These results highlight the need for formulae derived from reference samples from a wider range of body sizes to improve the reliability of existing methods. Am J Phys Anthropol, 2010. © 2009 Wiley‐Liss, Inc.     

Do big females have big pelves?

Previous studies have shown that maternal stature is a correlate of both pelvic size and reproductive efficiency. This study addresses the issue of body size and obstetric advantage. The relationship between pelvic size and three nonpelvic measures of body size is determined for females and males. The skeletal sample consists of blacks, whites, and Native Americans. The variables include 28 measures of the pelvis, length and head diameter of the femur, and clavicular length. The coefficient of multiple determination (CMD) is computed for each pelvic measure using multiple regression, with the three nonpelvic measures serving as the independent variables. Partial correlation coefficients are also calculated between each pelvic and nonpelvic variable, while controlling for the other two nonpelvic variables. The results show that all CMDs in females and all but one CMD in males are "low," i.e., below 33%. The sexes are nonsignificantly different in their CMDs for 22 of the 28 pelvic variables; of the six variables that are significantly different, five are of the midplane. The sexes are also broadly comparable in their partial correlations. The results are explained as follows. First, the concordance between the sexes in the relationship between pelvic size and nonpelvic measures of body size is due to their genetic similarity for homologous structures. Second, as pelvic size is at the minimum at the midplane, the sexual differences in CMDs are the result of selection with respect to obstetrics. Third, four explanations for the low CMDs are discussed: 1) lack of populationally or racially specific analysis; 2) nonlinear relationship between pelvic size and nonpelvic measures of body size; 3) combination of negative allometric selection between newborn body weight and maternal stature and weight with positive selection for maternal pelvic size; and 4) hormonally induced increase in pelvic capacity during parturition.     

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 and allometry in primate ossa coxae

Five measurements were taken on the ossa coxae of 454 adult primates representing Ceboidea, Cercopithecoidea and Hominoidea. Sex differences in these variables and their relationships to overall body size and sexual dimorphism were tested by means of Student's t-test and regression analysis. The study attempts to clarify the nature of primate pelvic sexual dimorphism, including allometric effects, and more specifically, test the assertion made by Mobb and Wood (1977) that sexual dimorphism in body size in not an important determinant in pelvic sex differences. Variables that contribute to the size of the birth canal tend to be larger in females than males in all taxa studied except two. In these, Hylobates and Alouatta, there were no significant differences between the sexes for any of the five variables. In general, sexual dimorphism in variables contributing to the size of the birth canal was correlated (r ? 0.8) with sexual dimorphism in body size. Furthermore, the coefficients of allometry underlying pelvic sex differences were shown to be moderately correlated (r ? 0.5) with sexual dimorphism in size. The influence of other adaptive factors on primate pelvic sexual dimorphism are also briefly discussed.     

Linking intra‐ and interspecific assortative mating: Consequences for asymmetric sexual isolation

Assortative mating is of interest because of its role in speciation and the maintenance of species boundaries. However, we know little about how within‐species assortment is related to interspecific sexual isolation. Most previous studies of assortative mating have focused on a single trait in males and females, rather than utilizing multivariate trait information. Here, we investigate how intraspecific assortative mating relates to sexual isolation in two sympatric and congeneric damselfly species (genus Calopteryx). We connect intraspecific assortment to interspecific sexual isolation by combining field observations, mate preference experiments, and enforced copulation experiments. Using canonical correlation analysis, we demonstrate multivariate intraspecific assortment for body size and body shape. Males of the smaller species mate more frequently with heterospecific females than males of the larger species, which showed less attraction to small heterospecific females. Field experiments suggest that sexual isolation asymmetry is caused by male preferences for large heterospecific females, rather than by mechanical isolation due to interspecific size differences or female preferences for large males. Male preferences for large females and male–male competition for high quality females can therefore counteract sexual isolation. This sexual isolation asymmetry indicates that sexual selection currently opposes a species boundary.     

Male size does not affect the strength of male mate choice for high-quality females in Drosophila melanogaster

Theory predicts that the strength of male mate choice should vary depending on male quality when higher-quality males receive greater fitness benefits from being choosy. This pattern extends to differences in male body size, with larger males often having stronger pre- and post-copulatory preferences than smaller males. We sought to determine whether large males and small males differ in the strength (or direction) of their preference for large, high-fecundity females using the fruit fly, Drosophila melanogaster. We measured male courtship preferences and mating duration to show that male body size had no impact on the strength of male mate choice; all males, regardless of their size, had equally strong preferences for large females. To understand the selective pressures shaping male mate choice in males of different sizes, we also measured the fitness benefits associated with preferring large females for both large and small males. Male body size did not affect the benefits that males received: large and small males were equally successful at mating with large females, received the same direct fitness benefits from mating with large females, and showed similar competitive fertilization success with large females. These findings provide insight into why the strength of male mate choice was not affected by male body size in this system. Our study highlights the importance of evaluating the benefits and costs of male mate choice across multiple males to predict when differences in male mate choice should occur.     

Sexual dimorphism in the pelvic midplane and its relationship to Neandertal reproductive patterns

The fragmentary nature of the fossil record has limited the analysis of the Neandertal pelvis to the superior pubic ramus and the pelvic inlet. From an obstetric viewpoint, the pelvic midplane or “plane of least dimensions,” defined by the distance between the ischial spines, must be considered in the analysis of hominid reproduction. We examined the relationship between BSD and weight in a mixed sex hospital population undergoing diagnostic computed tomography (CT) scans (41 females and 40 males). Because femoral head diameter squared (FH²) has been used as a proxy for weight in skeletal populations, it was also analyzed with respect to BSD and weight. Bivariate regression analysis of BSD with other body dimensions indicates the presence of significant sex differences. In females, but not in males, weight is a statistically significant predictor of BSD. FH² is an even better predictor of BSD in females while nonsignificant in males. Although weight and FH² are significantly correlated with BSD in females, FH² does not predict weight in females as well as it does in males. The positive correlation between skeletal frame size and BSD in females is indicative of an evolutionary pattern that must take into account the pressures of reproduction. Our results indicate that critical dimensions of the pelvis must increase as the maternal skeleton becomes larger. These results provide a context for the interpretation of the reproductive patterns of a relatively robust hominid population like the Neandertals. © 1996 Wiley-Liss, Inc.     

Smaller beetles are better scramble competitors at cooler temperatures

The role of temperature is central to both organic evolution and ecological processes. However, how temperature affects selection on body size is unknown. We tested whether small seed beetles (Stator limbatus) have an advantage over large beetles during scramble competition for mates, and whether this advantage varies with temperature. Within lines of beetles artificially selected to be large versus small, small males have a significant advantage over large males in scramble competition for females because the former takeoff more quickly and thus reach females before larger males. Selection favouring small male body size is significantly (and substantially) more intense at cooler temperatures. The adaptive significance of small male body size thus depends on ambient temperature.     

Paternal body size affects reproductive success in laboratory-held zebrafish (<Emphasis Type="Italic">Danio rerio</Emphasis>)

Across many fish species, large females tend to exhibit higher individual reproductive success due to elevated fecundity and the provisioning of better conditioned eggs and offspring compared to small females. By contrast, effects of paternal body size on reproductive success are less well understood. We disentangled the maternal- and paternal-size dependent effects on reproductive output and early life history in zebrafish (Danio rerio). In the laboratory, females and males from four size categories (small, medium-sized, large and very large) were allowed to spawn freely in a full factorial design with 10 replicates per size combination. As expected, larger females produced more eggs and better conditioned offspring compared to smaller females. Male body size further contributed to zebrafish reproductive success: offspring sired by large males exhibited higher hatching probability and these offspring also hatched earlier and larger than offspring fertilized by small males. However, the largest males experienced lower mating success and received fewer eggs than males of the smaller size classes. While male body size substantially affected reproductive success in zebrafish, it remained unclear whether and to what degree direct paternal effects (e.g., related to sperm quality) or indirect paternal effects stemming from differential allocation patterns by females were the mechanism behind our findings. Answering this question constitutes an important future research topic.     

Hunter-gatherer studies and human evolution: A very selective review

Objectives: Lower thoracic widths and curvatures track upper pelvic widths and iliac blades curvatures in hominins and other primates (torso integration hypothesis). However, recent studies suggest that sexual dimorphism could challenge this assumption in Homo sapiens. We test the torso integration hypothesis in two modern human populations, both considering and excluding the effect of sexual dimorphism. We further assess covariation patterns between different thoracic and pelvic levels, and we explore the allometric effects on torso shape variation. Material and Methods: A sex-balanced sample of 50 anatomically connected torsos (25 Mediterraneans, 25 Sub-Saharan Africans) was segmented from computed tomography scans. We compared the maximum medio-lateral width at seventh–ninth rib levels with pelvic bi-iliac breadth in males and females within both populations. We measured 1,030 (semi)landmarks on 3D torso models, and torso shape variation, mean size and shape comparisons, thoraco-pelvic covariation and allometric effects were quantified through 3D geometric morphometrics. Results: Females show narrow thoraces and wide pelves and males show wide thoraces and narrow pelves, although this trend is more evident in Mediterraneans than in Sub-Saharans. Equal thoracic and pelvic widths, depths and curvatures were found in absence of sexual dimorphism. The highest strength of covariation was found between the lowest rib levels and the ilia, and allometric analyses showed that smaller torsos were wider than larger torsos. Conclusions: This is the first study testing statistically the torso integration hypothesis in anatomically connected torsos. We propose a new and more complex torso integration model in H. sapiens with sexual dimorphism leading to different thoracic and pelvic widths and curvatures. These findings have important implications in hominin body shape reconstructions.     

Size Dependent Sexual Selection in Drosophila ananassae

Mate choice based on body size is widespread and can have numerous consequences. We present data, which show the effect of male and female body size on sexual selection in Drosophila ananassae. The relationships between wing size, locomotor activity, mating latency, courtship pattern, fertility and mating success were studied. Mating latency was negatively correlated with wing length and with locomotor activity, while wing length and locomotor activity was positively correlated in males as well as in females. In female- and male-choice, we found that mate choice influenced size-assortative mating by: (1) large and small males preferring to mate with large females, (2) large males successfully competing for large females, leaving small males to mate with small females. Males increased their reproductive success by mating with large and more fecund females. In addition, in pairs of long/short winged flies, long winged flies courted and mated more successfully than short winged flies and they also have longer duration of copulation and more progeny than short winged flies. We found sterile mating in pairs of small winged males and females.     

The relation between pelvic pressure and bladder pressure during pelvic perfusion with standardized flow rates

The aim of this study was to elucidate the relation between pelvic pressure and bladder pressure during pelvic perfusion with standardized flow rates. Anaesthetized Danish Landrace Breed pigs (n = 5) weighing 35-40 kg were studied. Transparenchymally two 6-F catheters were placed in both renal pelves for pressure measurement and perfusion. Transurethrally two catheters were placed in the bladder for pressure measurements and for urine collection and infusion. Bladder filling was done with a constant infusion rate of 45 ml/min during perfusion of both pelves with saline consecutively with the flow rates: 0, 2, 4, 6 and 8 ml/min during continuous measurement of bladder and bilateral pelvic pressure. The baseline diuresis varied from 0.4-1.0 ml/min. Without pelvic perfusion a negative pressure gradient between pelvis and bladder was seen demonstrating the importance of ureteral peristalsis. Pelvic perfusion with 2 ml/min showed that pelvic and bladder pressure were equal demonstrating weakening of ureteral peristalsis. During perfusion with higher flow rates pelvic pressure was higher than bladder pressure, showing that the positive gradient was important for urine transport. In conclusion ureteral peristalsis is important at low flow rates during increasing bladder pressure. At higher flow rates peristalsis weakens and the pressure gradient is the determining factor.     

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.     

Some evidence for different ecological pressures that constrain male and female body size

The dwarf morph of the Lake Tanganyika cichlid Telmatochromis temporalis uses empty snail shells as shelters and breeding sites in shell beds, in which many empty shells exist. Here, we assessed selection forces regulating body size in this fish. Field observations showed that large males tended to have a greater number of females in their territories, suggesting that sexual selection favours large males. Nonetheless, a transplant experiment suggested that male body size was limited by the ability to hide in empty shells from large piscivorous fish. In females, the number of ovarian eggs increased with body size, suggesting that fecundity selection favours large females. However, females are smaller than males. Females spawned eggs close to the apices inside the shells. The small space there would decrease the risk of egg predation by egg predators, and small body size of females may be a result of adaptation to spawn eggs in the small, safe spaces. This study provides support for the idea that male and female body sizes have been limited by different ecological pressures (predation on adult fish in males, predation on eggs in females), which has not been reported previously in any animal.