Skull shape asymmetry and the socioeconomic structure of an early medieval central european society

The socioeconomic structure of an early medieval society from the Mikul?ice settlement (Czech Republic) was studied on the basis of an evaluation of the fluctuating and directional asymmetry (DA) of skulls. Two distinct inhabited regions, castle and sub‐castle, were compared. Fluctuating asymmetry (FA) was used as a bioindicator of environmental stress, which is thought to have been different in the Mikul?ice castle and sub‐castle regions. DA is consistent with biomechanical loading, and it was expected to reflect different subsistence patterns. The material consisted of 129 crania from what was presumed to be a higher socioeconomic class (Mikul?ice castle) and 71 crania from middle and lower socioeconomic classes (Mikul?ice sub‐castle). As a comparative sample, 138 crania from modern, lower socioeconomic classes (Pachner Collection) were used. The three‐dimensional coordinates of 68 landmarks were digitized and analyzed using geometric morphometrics. In terms of DA, the highest values were recorded in the sub‐castle sample and confirmed their lower socioeconomic position, with a grittier and lower protein diet compared with the castle sample. In terms of FA, distinctive differences between the sexes were found. In males, no differences were observed between castle and sub‐castle, and the lowest FA values were recorded. In females, significantly higher values of FA were found, surprisingly in the castle sample, comparable with the more stressed Pachner Collection. We suspect that the FA reflects a more varied population of castle females as a consequence of patrilocality, although environmental stress remains a possibility. Am J Phys Anthropol, 2013. © 2013 Wiley Periodicals, Inc.     

Craniometrical variability and developmental stability. Two useful tools for assessing the population viability of Eurasian otter (Lutra lutra) populations in Europe.

Morphometrical univariate analyses of otter skulls collected over the past hundred years in European countries from presumed healthy populations were compared with skulls from presumed endangered populations. The average degree of sexual dimorphism of die European populations was found to be directly correlated to die skull size of die male otters. Fluctuating asymmetry (FA) in metric skull traits was analysed as an estimator of developmental stability. There was evidence for increased FA in different traits over time in some of the presumed endangered populations, and for a reduction in size of skull traits. In contrast, the healthy populations did not show any significant changes in the same traits during the same period. The reduced sexual dimorphism of the endangered populations is suggested to be a product of relaxed sexual selection and deteriorated habitat conditions. Environmental and genetic forces that may have shaped these patterns are discussed.     

Canalization and developmental instability of the fetal skull in a mouse model of maternal nutritional stress

Nutritional imbalance is one of the main sources of stress in both extant and extinct human populations. Restricted availability of nutrients is thought to disrupt the buffering mechanisms that contribute to developmental stability and canalization, resulting in increased levels of fluctuating asymmetry (FA) and phenotypic variance among individuals. However, the literature is contradictory in this regard. This study assesses the effect of prenatal nutritional stress on FA and among‐individual variance in cranial shape and size using a mouse model of maternal protein restriction. Two sets of landmark coordinates were digitized in three dimensions from skulls of control and protein restricted specimens at E17.5 and E18.5. We found that, by the end of gestation, maternal protein restriction resulted in a significant reduction of skull size. Fluctuating asymmetry in size and shape exceeded the amount of measurement error in all groups, but no significant differences in the magnitude of FA were found between treatments. Conversely, the pattern of shape asymmetry was affected by the environmental perturbation since the angles between the first eigenvectors extracted from the covariance matrix of shape asymmetric component of protein restricted and control groups were not significantly different from the expected for random vectors. In addition, among‐individual variance in cranial shape was significantly higher in the protein restricted than the control group at E18.5. Overall, the results obtained from a controlled experiment do not support the view of fluctuating asymmetry of cranial structures as a reliable index for inferring nutritional stress in human populations. Am J Phys Anthropol 154:544–553, 2014. © 2014 Wiley Periodicals, Inc.     

Fluctuating asymmetry in bank vole populations (Rodentia, Arvicolinae) reflects stress caused by landscape fragmentation in the Mont-Saint-Michel Bay

In intensively farmed, reclaimed areas (polders) of Mont-St-Michel Bay, France, bank voles ( Clethrionomys glareolus ) live in fragmented hedgerows, where populations are small and dispersal rates and genetic diversity are low. These small populations are likely to have been exposed to potential environmental and/or genetic stress. The sensitivity of development to stress can be measured by fluctuating asymmetry (FA). FA was calculated for three samples from a disturbed area and one sample from an adjacent, more connected and undisturbed landscape. Size FA was estimated from 16 measurements of the skull and teeth whilst shape asymmetry was estimated from the skull alone. Bank voles in fragmented hedgerows of the disturbed area had a higher degree of FA than bank voles from the more extensive and more connected hedges of the undisturbed area. These results were confirmed by the study of shape asymmetry, body mass and centroid size of the skull. There were no differences in FA between the three disturbed area samples. We conclude that FA does not reveal differences in the development of bank voles living in isolation under different local conditions in the various parts of the disturbed area. However, FA may allow differentiation between populations from greatly contrasting landscapes.  © 2003 The Linnean Society of London, Biological Journal of the Linnean Society , 2003, 80, 37–44.     

中国古人类颅容量的推算方法比较

古人类的体质特征和现代人不同,依据现代人头骨测量数据计算出来的公式往往不适于古人类颅容量的推算。获取古人类颅容量最准确的方法是复原出其内部的颅内模;然而,由于颅内模的复原工艺复杂,加上古人类头骨化石数量稀少且多数残破,如何准确地推算其颅容量,成为古人学者研究的难点问题之一。本文通过对中国境内发现的不同演化阶段的古人类颅容量推算方法的对比和验证,试图找出推算古人类颅容量的最适合的公式法。研究结果显示：1）早期现代人解剖特征同现代人基本接近,依据现代人头骨推算出来的回归方程可以用来推算其颅容量;2）直立人头骨厚重、脑颅低矮,体质特征不同于现代人,其颅容量的推算不能使用现代人公式法。依据本文中国直立人头骨测量数据推导出来的回程方程（C=-1301.944+60606L+0.718b+9.936h）适合其颅容量的推算。采用此直立人公式法,推算出蓝田直立人的颅容量为918 mL;3）古老型智人的体质特征位于直立人和现代人之间,对其颅容量的推算不能一概而论：体质特征接近直立人的,如大荔人、华龙洞6号,可采用直立人公式法;体质特征接近早期现代人的,如许昌1号,可采用现代人公式法;体质特征位于直立人和早期现代人中间位置的,如马坝人、金牛山人,其颅容量约等于采用现代人公式法和采用直立人公式法获得的颅容量的平均值。     

Environmental stress and fluctuating asymmetry in the grasshopper Chorthippus parallelus (Acrididae: Gomphocerinae)

We examined the effect of agrochemical pollution/fertilisation of the soil and climatic conditions at high altitudes on fluctuating asymmetry (FA) level in hindfemur and forewing lengths of the grasshopper Chorthippus parallelus (Zetterstedt). All traits and samples exhibited ideal FA. Forewings, which are less functionally significant in this species, generally exhibited greater FA than the more functionally significant femora. Forewing FA was also more affected by stress induced by climatic conditions at high altitudes than femur FA. The high altitude sample was the most asymmetric sample for both traits, followed by the two control samples. The two samples from agrochemically contaminated habitats showed the lowest asymmetry in femora and forewings of C. parallelus. These results suggest that high altitude imposes more stress on this grasshopper species than does agrochemical contamination although the stress response was more dramatic in the less functional trait.     

Strong nonlinear selection against fluctuating asymmetry in wild populations of a marine fish

Theoretical links between fluctuating asymmetry (FA) and fitness have led many to use FA as a proxy for average fitness. However, studies examining whether asymmetry actually correlates with individual fitness in wild populations are relatively rare and often use simple measures of association (e.g., correlation coefficients). Consequently, the pattern of selection on asymmetry in the wild is seldom clear. We examined selection on FA of pectoral fin morphology in two wild populations of a marine fish (the kelp perch; Brachyistius frenatus). As expected, variance in signed FA in each initial sample was significantly greater than that found in the surviving population, indicating selection against FA. Our estimate of the fitness surface confirmed perfect symmetry as the phenotypic optimum and indicated strong, nonlinear selection against asymmetry. No difference in the form of selection was detected between populations. However, the level of FA in the initial samples varied among populations, leading to an overall difference in the level of selective mortality. Our results suggest that selection on asymmetry in wild populations may be strongly nonlinear, and indicate that the demographic costs of asymmetry may play a substantial role in the dynamics of populations.     

Carabelli's trait in contemporary Slovenes and inhabitants of a medieval settlement (Sredisce by the Drava River)

The objectives of this study were to determine the total frequency, expression and asymmetry of Carabelli's trait in permanent dentitions of contemporary Slovenes and a medieval skeletal population from northeastern Slovenia. A total of 254 dental casts from contemporary Slovene children were examined. The population of a medieval settlement (10th-15th centuries), was represented by 94 skeletons. A modification of the method of Alvesalo and associates was used to classify Carabelli's trait on a five-grade scale. The trait was expressed on the upper first molars of 79.7% of the contemporary subjects and 75.8% of the medieval sample. Positive expressions of the trait were found in 10.1% of the contemporary subjects and 15.2% of the medieval sample. While the observed total frequency of the trait in both samples is characteristic of Europeans, the rates of positive expressions are surprisingly low but consistent with data from a recently published worldwide literature survey. Both populations showed a low rate of left-right fluctuating asymmetry of the trait. This finding might reflect a pronounced ability of individuals in the medieval population to buffer unfavourable influences from the environment and a relatively low level of environmental stress in the contemporary population.     

Morphological variability and developmental instability in subpopulations of the Eurasian badger (Meles meles) in Denmark

Aim Local populations from different geographical regions may differ in the selection regimes to which they are exposed. Differences in environmental factors and population density may affect the relative importance of different selective forces (e.g. natural vs. sexual selection). We suggest a direction of investigation concerned with the developmental instability of morphological traits. The goal is to disclose putative small‐scale geographical differences in the evolutionary forces, which may be hard to detect. Location Craniometrical investigations were carried out on ninety‐eight skulls and teeth of the Eurasian badger (Meles meles) collected during the period 1995–97 from three different populations in Denmark. One of these thrives at low population density, whereas the two others are characterized by high local density. Methods The skulls were investigated for developmental instability (DI) using fluctuating asymmetry (FA) as its estimator. FA was measured on canines, molars, premolar teeth and other skull and mandible traits. For the statistical analyses, we applied nonparametric permutation tests. Results Evidence was found suggesting differentiation among populations in mean degree of FA, and the FA values measured on canines were higher in the high‐density populations. FA of the canines was significantly higher in males than females, in contrast to FA of the other traits. Evidence of a negative relationship between canine size and their FA was found, whereas no significant correlations were found between the molar and premolar teeth measures and their FA. Main conclusions Our results suggest that canines could be under directional selection stemming from intrasexual competition, which may be stronger in high‐density zones. The other teeth investigated seem to be under a stabilizing regime hence their FA is mainly affected by environmental stresses. The negative relationship between canine size and FA found in males suggests the capacity of badgers to respond in an evolutionary way to environmental changes, despite the low genetic variability previously found at the molecular level.     

现代人头骨面部几项非测量性状的观察

用人类学观察法比较了中国云南、华北和欧洲现代人头骨的犬齿窝、颧骨缘突和梨状孔上外侧部膨隆的出现情况,以探讨其在人种、群体区分上是否有意义。结果：（1）在云南头骨,犬齿窝、颧骨缘突的出现率无性别差异,梨状孔上外侧部骨表面膨隆的出现率男性明显高于女性,而在欧洲头骨,这3项非测量必状的出现率男女间无差异。（2）上述3项性状,在中国云南和华北头骨（男性）的出现率无差异。（3）犬齿窝、梨状孔上外侧部骨表面膨隆的出现率,在现代欧洲人头骨明显高于现代中国人头骨。颧骨缘突的出现率在欧洲与中国现代人头骨上无差异。     

Fluctuating asymmetry and stress in a medieval Nubian population

Fluctuating asymmetry is commonly used as a bioindicator of developmental stress. This study addresses asymmetry under nutritional/systemic stress in the human craniofacial skeleton and its utility as an indicator of developmental instability. Crania from the diachronic Christian cemeteries at Kulubnarti (Sudanese Nubia) were chosen as a model for nutrition/systemic stress. Previous studies indicate that individuals from the Early Christian cemetery were subjected to greater developmental stress when compared with individuals from the Late Christian cemetery. Therefore, crania from the Early Christian cemetery should display a greater magnitude of fluctuating asymmetry than crania from the Late Christian cemetery. Thirty adult crania of comparable age and sex were selected from each population. Landmark coordinates were digitized in two separate trials and averaged to minimize error. Euclidean distance matrix analysis (EDMA) was used to measure and compare the magnitude of fluctuating asymmetry in each sample. Results indicate that crania from the Early Christian cemetery display greater amounts of fluctuating asymmetry than those from the Late Christian cemetery, as predicted. The degree of fluctuating asymmetry for each linear distance is highly correlated between the cemeteries, suggesting that all humans may share common patterns of fluctuating asymmetry in the skull. In contrast, there is little correlation between magnitude of fluctuating asymmetry and length of linear distance, between-subject variability, or measurement error. These results support the hypothesis that poor nutrition/systemic stress increases developmental instability in the human skull and that increased fluctuating asymmetry constitutes morphological evidence of this stress.     

FLUCTUATING ASYMMETRY IS NONGENETICALLY RELATED TO MATING SUCCESS IN DROSOPHILA BUZZATII

Abstract To date, there is still no consensus on the real significance of fluctuating asymmetry (FA) in evolutionary biology. Some studies have established links between FA and Darwinian fitness, and in a number of cases intermediate heritabilities for FA have been reported. However, many claims have been raised against the generality of these findings. I therefore tested if FA of a sexually selected trait (wing length) is indeed related to male mating success in Drosophila buzzatii from field and laboratory samples and whether FA has detectable heritability. Single, unsuccessful males had greater asymmetry for wing length than their mating counterparts both in nature and under nonoptimal rearing environments, but the higher FA in single males is most likely due to a poorer average phenotypic condition because there was no evidence of a genetic basis for this trait. Further evidence of an increase in FA under larval food stress is suggested when comparing the magnitude of the FA levels between stressful and optimal environments. On methodological grounds, a linear model is suggested that allows directional asymmetry (DA) and any genetic variation of DA that may be present to be statistically eliminated from estimates of FA.     

Pervasive genetic integration directs the evolution of human skull shape

It has long been unclear whether the different derived cranial traits of modern humans evolved independently in response to separate selection pressures or whether they resulted from the inherent morphological integration throughout the skull. In a novel approach to this issue, we combine evolutionary quantitative genetics and geometric morphometrics to analyze genetic and phenotypic integration in human skull shape. We measured human skulls in the ossuary of Hallstatt (Austria), which offer a unique opportunity because they are associated with genealogical data. Our results indicate pronounced covariation of traits throughout the skull. Separate simulations of selection for localized shape changes corresponding to some of the principal derived characters of modern human skulls produced outcomes that were similar to each other and involved a joint response in all of these traits. The data for both genetic and phenotypic shape variation were not consistent with the hypothesis that the face, cranial base, and cranial vault are completely independent modules but relatively strongly integrated structures. These results indicate pervasive integration in the human skull and suggest a reinterpretation of the selective scenario for human evolution where the origin of any one of the derived characters may have facilitated the evolution of the others.     

Absence of foramen spinosum and abnormal middle meningeal artery in cranial series

In comparative and evolutionary aspects in humans, the middle meningeal artery enters the cranium through the foramen spinosum, whereas in great apes the middle meningeal artery can enter the cranium through foramen spinosum, through foramen ovale or through petrosphenoid fissure. Generally, in nonhuman primates the anterior meningeal system is associated with the ophthalmic branch of the internal carotid artery. The vessels joining the two systems pass through the additional channels: the superior orbital fissure or through the cranio-orbital foramen. In anatomically modern humans, the absence of foramen spinosum involves abnormal development and course of the middle meningeal artery and it is usually accompanied with replacement of the conventional middle meningeal artery with such, arising from the ophthalmic artery system. In these cases the middle meningeal artery most often enters the middle cranial fossa through the superior orbital fissure and rarely through the meningo-orbital foramen. All skulls, investigated in the present study, belonged to adult individuals of both sexes, conditionally grouped into three cranial series--contemporary male, medieval male, and medieval female series. The absence of foramen spinosum was established only among the medieval male and female series--in 1 (0.70%) male and in 1 (0.72%) female skull on the right side and in 3 (2.13%) female skulls on the left side. In 1 (0.72%) female skull, a small atypically located foramen spinosum was established on the right side. In all of the described cases, the intracranial meningeal grooves started from the lateral edge of the superior orbital fissure and probably reflect the ophthalmic origin of the middle meningeal artery.     

Adaptations to humeral torsion in medieval Britain

Adaptations to the humeral torsion angle have been identified in the professional throwing athlete. This movement pattern increases the humeral torsion angle, and also increases the extent of external rotation movements in the dominant, throwing limb when compared with the nondominant limb. The purpose of this paper is to test the hypothesis that the humeral torsion angle is an adaptation to upper limb use. This project examines the humeral torsion angle in a number of medieval British populations, as well as a modern cadaver-based sample. The results identify significant differences in the humeral torsion angle both between and within male (P < 0.001, ANOVA) and female (P < 0.014, ANOVA) populations, although the results are not consistent with expected behavior patterns. Statistically significant differences between males and females within the same site were identified in 2 of the 5 samples examined. The mean level of bilateral asymmetry does not approach that reported for the professional throwing athlete. However, a number of individuals have high levels of asymmetry in excess of that identified in the professional throwing athlete. This analysis demonstrates the need for individual rather than population-based analyses, as the heterogeneity within population samples obscures individual variation in activity patterns. The diversity within British medieval society and a lack of specific known behaviors prevent further identification of the functional significance of the humeral torsion angle within the archaeological record examined here.     

Fluctuating asymmetry of craniometric characters in rodents (Mammalia: Rodentia): interspecific and interpopulational comparisons

Ontogenetic instability was studied in four rodent species (Ellobius talpinus, Microtus arvalis, M. rossiaemeridionalis, Mus musculus). It was measured by fluctuating asymmetry (FA) of 21 craniometric characters. Each species was represented by two populations differing in average level of chromosomic instability and in degree of anthropogenic influence. Relation between FA and individual age was observed in none of the studied species. The level of cranial structure FA in rodents is probably formed during embryogeny or early postnatal ontogeny and is not changing significantly during the further development. The studied species showed distinct differences in the degree of ontogenetic instability of the axial skull, but not of the mandible. No connection was found between craniometric character FA and technogenic stress. The differences of cranial FA level among the studied species of rodents agree with their systematic position and the sequence of their evolutionary divergence.     

Adolescent health in medieval Serbia: signs of infectious diseases and risk of trauma

Although pattern of health in adults has been frequently assessed in past human populations, health status of adolescents as a distinct life stage has usually been overlooked. Inconsistency in number and meaning of recognised age categories in anthropological literature, as well as chronological age ranges used to define them, further complicate the interpretation of adolescent health. In this study, we analysed signs of pathological conditions on skeletal remains of 81 adolescents from a medieval site of Stara Torina (northern Serbia). Diagnostic palaeopathological procedures comprised gross examination, digital radiography, and histological analysis.Skeletal signs of anaemia such as cribra orbitalia and other porotic phenomena as well as signs of non-specific bone infection were observed frequently, while evidence of bone trauma was recorded in a very low percentage of individuals. In addition, we recorded two conditions relatively rarely observed in palaeopathological contexts: a case of skull and vertebral asymmetry indicative of congenital muscular torticollis, and a case of a fibrous cortical defect on distal femur. Comparison with available information from other medieval adolescent samples from Serbia demonstrated that while mortality was relatively constant throughout the sample, Stara Torina showed a much higher occurrence of bone disease. Characteristics of observed skeletal conditions, supported by available historical reports, suggest that the health of medieval adolescents in the examined population was most significantly affected by infectious processes.     

A measure of asymmetry of the skulls of odontocete whales

Linear measurements were made of the asymmetry of the skull which is characteristic of odontocete whales. Of the living genera (except Physeter, Tasmacetus, Stenella, Feresa and Lissodelphis) at least one specimen was measured; 317 skulls in all. One species, Stenodelphis blainvillei , proved symmetrical. All other species had leftward deviation of the nasal prominence. No specimen showed inverse asymmetry.
When, for species or families, regressions were formed of the asymmetry measure on skull length the slopes formed a sequence such that Kogia >monodontids>delphinoids (except Orcinus)>ziphiids> Orcinus. In some species of delphinoid, in the delphinoids as a group, and more strongly in Monodon , the proportion of asymmetry to skull length increased with increasing size. In the ziphiids, as a family, asymmetry increased roughly linearly with bodily size. Orcinus orca proved an anomalous delphinoid in having little asymmetry for its size. The asymmetry of the skull of Monodon monoceros is independent of the possession of a tusk.
It is suggested that the leftward deviation of the dorsal aspect of the skull acts to conserve the symmetry of the body surface by countering an opposite asymmetry of the upper breathing passages, an asymmetry which is primary but whose function is unknown.     

Unilateral and bilateral expression of a quantitative trait: asymmetry and symmetry in coronal craniosynostosis

Bilateral symmetry in vertebrates is imperfect and mild asymmetries are found in normal growth and development. However, abnormal development is often characterized by strong asymmetries. Coronal craniosynostosis, defined here as consisting of premature suture closure and a characteristic skull shape, is a complex trait. The premature fusion of the coronal suture can occur unilaterally associated with skull asymmetry (anterior plagiocephaly) or bilaterally associated with a symmetric but brachycephalic skull. We investigated the relationship between coronal craniosynostosis and skull bilateral symmetry. Three-dimensional landmark coordinates were recorded on preoperative computed tomography images of children diagnosed with coronal nonsyndromic craniosynostosis (N = 40) and that of unaffected individuals (N = 20) and analyzed by geometric morphometrics. Our results showed that the fusion pattern of the coronal suture is similar across individuals and types of coronal craniosynostosis. Shape analysis showed that skulls of bilateral coronal craniosynostosis (BCS) and unaffected individuals display low degrees of asymmetry, whereas right and left unilateral coronal craniosynostosis (UCS) skulls are asymmetric and mirror images of one another. When premature fusion of the coronal suture (without taking into account cranial dysmorphology) is scored as a qualitative trait, the expected relationship between trait frequency and trait unilateral expression (i.e. negative correlation) is confirmed. Overall, we interpret our results as evidence that the same biological processes operate on the two sides in BCS skulls and on the affected side in UCS skulls, and that coronal craniosynostosis is a quantitative trait exhibiting a phenotypic continuum with BCS displaying more intense shape changes than UCS.