Obstetric adaptation of the human pelvic inlet based on a sample of females and males matched for body size, and comparison with three species of primates that are sexually monomorphic in body size.

Human males are larger than females in body size; females are larger than males in pelvic size. Percentage pelvic sexual size dimorphism (sex with larger mean/sex with lower mean*100%) is used to infer selection for obstetric adaptation. As pelvic size is associated with body size, interpretation of percentage sexual size dimorphism necessitates control of body size. This study uses a larger sample of 221 females and 204 males to identify a subsample of females and males matched for femoral length, femoral head diameter, and clavicular length (25 females, 46 males). For five measures of the pelvic inlet, the human sample with control of body size is compared with three primate species that are sexually monomorphic in body size and give birth to newborns with higher relative body masses than those of humans: Aotus azarae (15 females, 19 males), Hylobates lar (29 females, 29 males), and Saguinus geoffroyi (28 females, 30 males). Results show that humans (112.0%) have the highest percentage sexual size dimorphism of pelvic inlet circumference: A. azarae (108.2%), H. lar (105.4%), and S. geoffroyi (103.3%). Humans have higher sexual size dimorphism of inlet circumference despite lower relative newborn body mass compared to the other species because human newborns are of larger body mass than predicted for an anthropoid with our adult female body mass. In humans, birthing of bigger babies than predicted for our body mass increases selection pressure on females for enlargement of the pelvic inlet.

Rudimentary, "functionless" first metapodials of Canis latrans: Variation and association in length with longer, functional metapodials.

A tenet of evolutionary theory is that phenotypic variation of a trait is inversely related to the intensity of stabilizing selection pressure. Among homologous bones, such as metapodials, a rudimentary, "nonfunctional" bone is expected to be more variable in length than nonrudimentary bones. This study compares variation and association in length among metapodials using 277 adult skeletons of Canis latrans. Canis latrans has a short, "functionless" first metacarpal (mc1) and "rudimentary, vestigial" first metatarsal (mt1). Results show that among the 10 metapodials, mt1 has the highest variation in length; other metapodials do not differ significantly from one another in their variation. Correlation coefficients for length of mc1 and mt1 with their ipsilateral metapodials 2-5 are significantly lower than coefficients for all other ipsilateral pairs. The correlation coefficient between left and right mt1 is significantly the lowest among all bilateral pairs of metapodials. Results are interpreted as follows. Mt1's high variation and low association in length are the outcome of less intense stabilizing selection pressure compared with other metapodials. The nonsignificant difference for variation in length between mc1 and metapodials 2-5 may be that mc1 is functional for development of a pollical dewclaw that helps restrain small prey.

Proximate cause, anatomical correlates, and obstetrical implication of a supernumerary lumbar vertebra in humans.

OBJECTIVES: Three issues are considered on variation in number of presacral vertebrae (PSV) in humans: (1) sexual difference in number of PSV, (2) inactivation of Hoxd-11 gene as etiology for a supernumerary lumbar vertebra, and (3) anatomical correlates of a supernumerary lumbar vertebra, including lumbar-sacral nearthrosis, and pelvic size. MATERIALS AND METHODS: Sample was 407 skeletonized females and 1,318 males from United States; ages at death were 20 to 49 years. Two subsamples of males were used: (1) 98 with modal numbers of cervical, thoracic, lumbar, and sacral vertebrae (PSV = 24) and (2) 45 with a supernumerary lumbar vertebra but modal numbers for other vertebral segments (PSV = 25). Measurements were taken of ulna, second metacarpal, vertebrae, femur, and pelvis; presence of lumbar-sacral nearthrosis was observed. RESULTS: Although 90% of females and males have 24 PSV, females have higher frequency of 23 PSV and males have higher frequency of 25 PSV. Compared to males with 24 PSV, males with 25 PSV and supernumerary lumbar vertebra show (1) no difference in anatomies associated with inactivation of Hoxd-11, and (2) higher frequency of lumbar-sacral nearthrosis and smaller pelvic inlet circumference. DISCUSSION: Sexual difference in number of PSV may be due to tempo of somite formation and Hox gene activation. Hypothesis is not supported that a supernumerary lumbar vertebra is due to inactivation of Hoxd-11. The presence of a supernumerary lumbar vertebra is associated with small pelvic inlet circumference, which can be obstetrically disadvantageous.

Sacral Variability in Tailless Species: Homo sapiens and Ochotona princeps.

Homo sapiens is variable in number of sacral vertebrae, and this variability can lead to obstetrical complication. This study uses the comparative method to test the hypothesis that sacral variability in H. sapiens is associated with absence of a tail. Three species of lagomorphs are studied: Ochotona princeps (N = 271), which is tailless, and Lepus californicus (N = 212) and Sylvilagus audubonii (N = 206), which have tails. Results show that O. princeps has (1) higher diversity index for number of sacral vertebrae (0.49) compared to L. californicus (0.25) and S. audubonii (0.26) and (2) significantly higher percentage of individuals with the species-specific nonmodal number of sacral vertebrae (43.9%) compared to L. californicus (14.2%) and S. audubonii (15.5%). Comparison of H. sapiens (N = 1,030; individuals of age 20-39 years) with O. princeps shows similarities between the species in diversity index (also 0.49 in H. sapiens) and percentage of individuals with nonmodal number of sacral vertebrae (37.3% in H. sapiens). Homeotic transformation best explains the results. H. sapiens and O. princeps show propensity for caudal shift at the sacral-caudal border (i.e., homeotic transformation of the first caudal vertebra to a sacral vertebra). Caudal and cranial shift among presacral vertebrae increases or decreases this propensity, respectively. Increase in number of sacral vertebrae in H. sapiens by homeotic transformation reduces pelvic outlet capacity and can be obstetrically hazardous. Anat Rec, 300:798-809, 2017. © 2017 Wiley Periodicals, Inc.

Pelvic sexual dimorphism among species monomorphic in body size: relationship to relative newborn body mass.

Females have larger pelves than males among eutherians to mitigate obstetrical difficulty. This study addresses 3 issues concerning pelvic sexual dimorphism using 8 species that are sexually monomorphic in nonpelvic size: Aotus azarae , Castor canadensis , Dasypus novemcinctus , Hylobates lar , Saguinus geoffroyi , Sciurus carolinensis , Sylvilagus floridanus , and Urocyon cinereoargenteus . Using published data to compute the index of relative newborn body mass (RNBM = [newborn body mass/adult female body mass]100%) for 266 eutherian species, A. azarae , H. lar , and S. geoffroyi are characterized as giving birth to relatively large newborns and the other 5 species as giving birth to relatively small newborns. The 3 issues are, compared to species giving birth to relatively small newborns, whether species that give birth to relatively large newborns have 1) higher magnitude of pelvic sexual size dimorphism (SSD), 2) lower prevalence of pelvic joint fusion, and 3) dissociation between pelvic and nonpelvic sizes. Nine measures of the pelvis were taken, and fusion of interpubic and sacroiliac joints was observed. Species grouped by high and low RNBM do not differ significantly in magnitude of SSD of pelvic inlet circumference. Species with high RNBM have significantly lower prevalence of interpubic joint fusion than those with low RNBM. Sexes do not differ in their multiple correlation coefficients between inlet circumference and nonpelvic body size in 7 of 8 species. Results suggest that 1) there are multiple anatomical pathways for pelvic obstetrical sufficiency, 2) an unfused interpubic joint is obstetrically advantageous, and 3) relative newborn size does not change the association between pelvic and nonpelvic size in females and males.

Fusion of coccyx to sacrum in humans: prevalence, correlates, and effect on pelvic size, with obstetrical and evolutionary implications.

Humans do not have a tail, but we have four rudimentary coccygeal vertebrae. This study considers several issues pertaining to fusion of the coccyx to the sacrum, including prevalence, sexual differences, effect on pelvic size, and obstetrical and evolutionary implications. Previous research on sacral-coccygeal fusion has reported: (1) lower prevalence in females than males, (2) prevalence increases with age, (3) range in prevalence among 13 samples from 0 to 72%, and (4) obstetrical complications. This study uses a sample of 2,354 American skeletons of known sex, age 20 years and older to ascertain prevalence of sacral-coccygeal fusion and to evaluate some of its correlates. Results show that the sexes do not differ in prevalence of sacral-coccygeal fusion for five of seven decades of life, but that prevalence does increase with advancing age-from 24 to 47% from the third to eighth decades of life in females. Pelvimetric analysis of 132 females shows that those with sacral-coccygeal fusion have a shorter posterior sagittal diameter of the outlet compared to those without fusion; more than half of those with sacral-coccygeal fusion have an obstetrically contracted posterior sagittal diameter. Shortening of the posterior sagittal diameter is important, because its conjoint occurrence with a narrow subpubic arch may result in an obstetrically inadequate outlet. This study concludes that sacral-coccygeal fusion is a principal contributor to the evolution of sexual dimorphism in sacral angulation, which is a determinant of the length of the posterior sagittal diameter of the outlet.

Sacralization is not associated with elongated cervical costal process and cervical rib.

Cervical rib/elongated costal process of the seventh cervical vertebra and sacralization of a lumbar vertebra are associated with clinical problems-neurological, vascular, and obstetrical. A previous study reported an association between these morphologies, and suggested that clinicians use presence of one trait to predict presence of the other. This study tested three hypotheses from this association: costal process length among individuals with sacralization differs from that among individuals without sacralization for: (1) only the seventh cervical vertebra, (2) only transitional presacral vertebrae-seventh cervical, twelfth thoracic, and fifth lumbar, and (3) presacral vertebrae in general. Skeletons of 961 individuals between ages 20 and 49 years from the United States were surveyed for sacralization. Costal process length was measured on 100 individuals with sacralization and 184 without sacralization for cervical vertebrae 3 to 7, thoracic vertebrae 11 and 12, and all lumbar vertebrae. Cervical rib was evaluated for 102 individuals with sacralization and 472 without sacralization. Results showed that 11% (105 of 961) of individuals have sacralization. Compared to individuals without sacralization, those with sacralization: (1) have significantly longer costal process for the last lumbar vertebra, but are nonsignificantly different for costal process lengths of other vertebrae, (2) are nonsignificantly different in prevalence of cervical rib-2.9% with sacralization and 0.4% without sacralization, and (3) are significantly more likely to have an extra presacral vertebra. Clinically, results suggest that sacralization is not a predicate for either cervical rib or elongated costal process of the seventh cervical vertebra.

High assimilation of the sacrum in a sample of American skeletons: prevalence, pelvic size, and obstetrical and evolutionary implications.

High assimilation sacrum is fusion of the caudal-most lumbar vertebra to the first sacral vertebra. Previous studies have shown that high assimilation is associated with clinical problems, including obstetrical difficulty. This study used adult American males (n = 1,048) and females (n = 1,038) of the Hamann-Todd and Terry skeletal collections to determine the prevalence of high assimilation and its effect on pelvic size, and to consider the obstetrical and evolutionary implications of high assimilation. The prevalence of high assimilation in this sample is 6.3%, with males and females not differing significantly from one another in their prevalence. This prevalence is near the median for that reported in 41 other samples. In both males and females, individuals with high assimilation have significantly longer anteroposterior and posterior sagittal diameters of the inlet, and shorter sacrum compared to those with a nonassimilated sacrum. Females with high assimilation have a significantly narrower sacral angulation (i.e., reduced inclination of ventral axis of sacrum), and shorter posterior sagittal diameter of the outlet compared to those with a nonassimilated sacrum. A short posterior sagittal diameter of the outlet is associated with childbirth difficulty. As high assimilation is partial homeotic transformation of a lumbar vertebra, this study supports previous research that homeotic transformation of vertebrae is selectively disadvantageous.

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.

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.

Variability of metapodials in primates with rudimentary digits: Ateles geoffroyi, Colobus guereza, and Perodicticus potto.

A tenet of evolutionary theory is that, within a species, phenotypic variability is inversely related to the intensity of stabilizing selection. A corollary is that a rudimentary or vestigial structure should be highly variable. This relationship between rudimentation and variability, however, may simply be part of a continuum, as several studies have shown that variability and size of a structure are inversely related. This study tests whether the first metacarpal (MC1) in Ateles geoffroyi and Colobus guereza and the second metacarpal (MC2) in Perodicticus potto are highly variable in their lengths relative to their other metapodials. The former two species have rudimentary thumbs, and the latter species has a rudimentary index finger. Fourteen other species of primates are included in the comparison. The results show that MC1 in A. geoffroyi and C. guereza and MC2 in P. potto are the relatively shortest first and second metapodials, respectively, in this sample of primates. However, an intraspecific analysis shows that neither MC1 in A. geoffroyi and C. guereza nor MC2 in P. potto is significantly more variable than the other metapodials. Nevertheless, an interspecific analysis shows that MC1 in A. geoffroyi and C. guereza is relatively the most variable among the first metapodials (i.e., MC1 and first metatarsal) in this study. MC2 in P. potto, however, is of relatively low variability compared with the other primates. These contrasting results are interpreted in terms of the developmental and evolutionary biology of digits.

VARIABILITY OF A VESTIGIAL STRUCTURE: FIRST METACARPAL IN COLOBUS GUEREZA AND ATELES GEOFFROYI.

A tenet of evolutionary theory is that, under conditions of stabilizing selection, phenotypic variability is inversely related to selection intensity. Correspondingly, a nonfunctional, vestigial structure is expected to be highly variable relative to its functional homologue. This study tests the hypothesis that species with a vestigial pollex, Colobus guereza and Ateles geoffroyi, have a first metacarpal whose length is both highly variable relative to, and poorly correlated with, the lengths of the other metapodials. The results are consistent with the hypothesis, though this combination of traits is also found in Presbytis rubicunda and Presbytis cristata. The latter two species have functional, albeit miniature, pollices. This study also demonstrates a general, inverse relationship among anthropoid primates between relative length of the first metapodial and its relative variability. These results suggest that elevated variability accompanies structural reduction.