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.     

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.     

There is an obstetrical dilemma: Misconceptions about the evolution of human childbirth and pelvic form

Compared to other primates, modern humans face high rates of maternal and neonatal morbidity and mortality during childbirth. Since the early 20th century, this “difficulty” of human parturition has prompted numerous evolutionary explanations, typically assuming antagonistic selective forces acting on maternal and fetal traits, which has been termed the “obstetrical dilemma.” Recently, there has been a growing tendency among some anthropologists to question the difficulty of human childbirth and its evolutionary origin in an antagonistic selective regime. Partly, this stems from the motivation to combat increasing pathologization and overmedicalization of childbirth in industrialized countries. Some authors have argued that there is no obstetrical dilemma at all, and that the difficulty of childbirth mainly results from modern lifestyles and inappropriate and patriarchal obstetric practices. The failure of some studies to identify biomechanical and metabolic constraints on pelvic dimensions is sometimes interpreted as empirical support for discarding an obstetrical dilemma. Here we explain why these points are important but do not invalidate evolutionary explanations of human childbirth. We present robust empirical evidence and solid evolutionary theory supporting an obstetrical dilemma, yet one that is much more complex than originally conceived in the 20th century. We argue that evolutionary research does not hinder appropriate midwifery and obstetric care, nor does it promote negative views of female bodies. Understanding the evolutionary entanglement of biological and sociocultural factors underlying human childbirth can help us to understand individual variation in the risk factors of obstructed labor, and thus can contribute to more individualized maternal care.     

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.     

Effects of parity on pelvic size and shape dimorphism in Mus

The pelvis is a sexually dimorphic structure and although the causes of that dimorphism have long been studied, relatively little is known regarding the effects of partuitive events on the magnitude of that dimorphism. Here, we use a sample of Mus musculus domesticus to contrast dimorphism in body length and os coxae size and shape between males and parous and nulliparous females. We also test for correlations between relative litter size (L/M) and relative offspring size (O/M) with body length and os coxae size and shape in parous females. Males had greater body length than nulliparous females but were not different from parous females. Females as a whole had the largest os coxae, with parous females having the largest and males the smallest. Os coxae shape was also significantly different between groups and was most divergent between parous females and males than between nulliparous females and males. Os coxae shape differences between females are associated with differences in body length between females and O/M is correlated with os coxae shape in parous females such that females with the largest offspring have the most divergent shapes along the relative warp one axis. Pelvic shape differences between males and females were consistent with previous findings in other taxa which identify the pubo‐ischial complex as the primary region of dimorphism. J. Morphol. 2009. © 2009 Wiley‐Liss, Inc.     

Spondylolysis of the sacrum in Alaskan and Canadian Inuit skeletons

Spondylolysis of the lower back, particularly that involving the isthmus between the superior and inferior articular processes (pars inter-articularis), is generally attributed to stress fracturing caused by movement of the affected vertebra relative to the vertebra below. The finding of isthmic spondylolysis in the first vertebra of a fused sacrum is thus unusual and requires explanation. Although unrepresented in the clinical literature, sacral spondylolysis has been reported for archaeological specimens and appears to be especially prevalent in North American Inuit. A study of 373 Inuit sacra from Alaska and Canada produced 16 examples of spondylolysis (eight from each area). All but one of the affected individuals were male, and nearly all were young adults, many between 18 and 20 years of age. All cases of sacral spondylolysis observed in this study were judged to have resulted from stress fracturing that occurred while S1 was still unfused, and most appear to have been in the process of healing, following fusion of S1 with S2, when death occurred. The high frequency observed in these people is attributed to unusual stresses becoming concentrated in the lower back of adolescent males due to such activities as weight lifting, wrestling, kayak paddling, and harpooning, combined with, and perhaps even contributing to, delayed maturation (S1-S2 fusion) of the sacrum. © 1996 Wiley-Liss, Inc.     

Biomechanics of the hip and birth in early Homo

A complex of traits in the femur and pelvis of Homo ereclus and early “erectus-like” specimens has been described, but never satisfactorily explained. Here the functional relationships between pelvic and femoral structure in humans are explored using both theoretical biomechanical models and empirical tests within modern samples of diverse body form (Pecos Amerindians, East Africans). Results indicate that a long femoral neck increases mediolateral bending of the femoral diaphysis and decreases gluteal abductor and hip joint reaction forces. Increasing biacetabular breadth along with femoral neck length further increases M-L bending of the femoral shaft and maintains abductor and joint reaction forces at near “normal” levels. When compared to modern humans, Homo erectus and early “erectus-like” specimens are characterized by a long femoral neck and greatly increased M-L relative to A-P bending strength of the femoral shaft, coupled with no decrease in hip joint size and a probable increase in abductor force relative to body size. All of this strongly suggests that biacetabular breadth as well as femoral neck length was relatively large in early Homo. Several features preserved in early Homo partial hip bones also indicate that the true (lower) pelvis was very M-L broad, as well as A-P narrow. This is similar to the lower pelvic shape of australopithecines and suggests that nonrotational birth, in which the newborn's head is oriented transversely through the pelvic outlet, characterized early Homo as well as Australopithecus. Because M-L breadth of the pelvis is constrained by other factors, this may have limited increases in cranial capacity within Homo until rotational birth was established during the late Middle Pleistocene. During or after the transition to rotational birth biacetabular breadth decreased, reducing the body weight moment arm about the hip and allowing femoral neck length (abductor moment arm) to also decrease, both of which reduced M-L bending of the proximal femoral shaft. Variation in femoral structural properties within early Homo and other East African Early Pleistocene specimens has several taxonomic and phylogenetic implications. © 1995 Wiley-Liss, Inc.     

Intermarriage and Jewish identity: The implications for pluralism and assimilation in American society*

Born across racial lines, Cedric Dover and Barack Obama both came to identify with the African American community. By contrasting the lives and ideas of two mixed-race individuals, one born in Calcutta and the other in Hawaii, this article examines cosmopolitanism, racial formation and the promise of the ‘post-racial’. A ‘Eurasian’ intellectual born in Calcutta in 1904, Dover developed a coloured cosmopolitanism that mirrors in revealing ways Obama's approach to race. Both men embraced blackness while transcending the boundaries of race and nation. Dover and Obama developed a conception of race as freedom – not freedom from race or of a particular race, but the freedom to embrace race without sacrificing other affiliations.     

Evolution of growth habit, inflorescence architecture, flower size, and fruit type in Rubiaceae: its ecological and evolutionary implications

During angiosperm evolution, innovations in vegetative and reproductive organs have resulted in tremendous morphological diversity, which has played a crucial role in the ecological success of flowering plants. Morindeae (Rubiaceae) display considerable diversity in growth form, inflorescence architecture, flower size, and fruit type. Lianescent habit, head inflorescence, small flower, and multiple fruit are the predominant states, but arborescent habit, non-headed inflorescence, large flower, and simple fruit states occur in various genera. This makes Morindeae an ideal model for exploring the evolutionary appearances and transitions between the states of these characters. We reconstructed ancestral states for these four traits using a bayesian approach and combined nuclear/chloroplast data for 61 Morindeae species. The aim was to test three hypotheses: 1) self-supporting habit is generally ancestral in clades comprising both lianescent and arborescent species; 2) changes from lianescent to arborescent habit are uncommon due to "a high degree of specialization and developmental burden"; 3) head inflorescences and multiple fruits in Morindeae evolved from non-headed inflorescences and simple fruits, respectively. Lianescent habit, head inflorescence, large flower, and multiple fruit are inferred for Morindeae, making arborescent habit, non-headed inflorescence, small flower, and simple fruit derived within the tribe. The rate of change from lianescent to arborescent habit is much higher than the reverse change. Therefore, evolutionary changes between lianescent and arborescent forms can be reversible, and their frequency and trends vary between groups. Moreover, these changes are partly attributed to a scarcity of host trees for climbing plants in more open habitats. Changes from large to small flowers might have been driven by shifts to pollinators with progressively shorter proboscis, which are associated with shifts in breeding systems towards dioecy. A single origin of dioecy from hermaphroditism is supported. Finally, we report evolutionary changes from headed to non-headed inflorescences and multiple to simple fruits.     

Genome size in 21 Artemisia L. species (Asteraceae, Anthemideae): systematic, evolutionary, and ecological implications.

Genome size was estimated by flow cytometry in 24 populations belonging to 22 Artemisia taxa (21 species, 1 with two subspecies), which represent the distinct subgenera, life forms, basic chromosome numbers, and ploidy levels in the genus. 2C nuclear DNA content values range from 3.5 to 25.65 pg, which represents a more than sevenfold variation. DNA content per haploid genome ranges from 1.75 to 5.76 pg. DNA amount is very well correlated with karyotype length and ploidy level. Some variations in genome size have systematic and evolutionary implications, whereas others are linked to ecological selection pressures.     

High genetic divergence of hormogastrid earthworms (Annelida, Oligochaeta) in the central Iberian Peninsula: evolutionary and demographic implications

Hormogastridae earthworms are highly important for the functioning of the Mediterranean soil system. However, little is known about the species distribution and genetic diversity of these soil invertebrates. In the present study, the genetic differentiation and gene flow were studied among populations of hormogastrids from the central Iberian Peninsula. A 648-bp portion of the mitochondrial cytochrome c oxidase I gene was sequenced for 82 individuals from 7 localities, resulting in the identification of 38 haplotypes exclusive to localities. All of the individuals were morphologically identified as Hormogaster elisae , but the high genetic divergence found among populations (up to 20.20%) suggests the occurrence of more than one cryptic species within this region. Further analysis of the phylogenetic relationships revealed six different evolutionary lineages coincident with geographical location, including the two nearest populations Molar and Redueña as one evolutionary unit. From these results, at least three new species could be inferred, in addition to the morphospecies H. elisae s.s . Partitioning of genetic variance among populations indicated that isolation by distance was the primary agent for differentiation of the investigated hormogastrid populations. Our data suggest that the evolutionary lineages for H. elisae s.l. originated between the late Miocene and the early Pleistocene, but that mtDNA genealogies coalesce on a more recent scale of a few thousand years.     

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.     

Conflicting selection pressures on seed size: evolutionary ecology of fruit size in a bird-dispersed tree, Olea europaea

Recent evidence indicates that fruit size has evolved according to dispersers' size. This is hypothesized to result from a balance between factors favouring large seeds and dispersers setting the maximum fruit size. This hypothesis assumes that (1) the size of fruits that can be consumed by dispersers is limited, (2) fruit and seed size are positively correlated, and (3) the result of multiple selection pressures on seed size is positive. Our studies on the seed dispersal mutualism of Olea europaea have supported the first and second assumptions, but valid tests of the third assumption are still lacking. Here we confirm the third assumption. Using multiplicative fitness components, we show that conflicting selection pressures on seed size during and after dispersal reverse the negative pattern of selection exerted by dispersers.