Sex determination from the occipital condyle: discriminant function analysis in an eighteenth and nineteenth century British sample

Fragmentary human remains compromised by different types of inhumation, or physical insults such as explosions, fires, and mutilations may frustrate the use of traditional morphognostic sex determination methods. The basicranium is protected by a large soft tissue mass comprising muscle, tendon, and ligaments. As such, the occipital region may prove useful for sex identification in cases of significantly fragmented remains. The aims of this paper are to (1) evaluate sexual dimorphism in British cranial bases by manually recorded unilateral and bilateral condylar length and width as well as intercondylar measurements and (2) develop discriminant functions for sex determination for this cranial sample. The crania selected for this study are part of the 18th-19th century documented skeletal collection of St. Bride's Church, Fleet Street, London. Adult human skulls (n = 146; male75/female71) were measured to derive statistical functions. Results indicated that expression of sexual dimorphism in the occipital condylar region within the St. Bride's population is demonstrable but low. Crossvalidated classification accuracy ranged between 69.2 and 76.7%, and sex bias ranged from 0.3 to 9.7%. Therefore, the use of discriminant functions derived from occipital condyles, especially in British skeletal populations, should only be considered in cases of fragmented cranial bases when no other morphognostic or morphometric method can be utilized for sex determination.     

现代人头骨断面轮廓的性别鉴定——基于几何形态测量的研究

人类头骨在大小、粗壮度和形状上具有性别差异。长期以来,性别鉴定的方法主要是根据头骨的大小和粗壮度,用肉眼进行粗略观察。受研究方法的限制,忽略了头骨细微的解剖结构及形状差别。为获取头骨性别差异的更多信息,探究头骨断面轮廓的性别差异,本文选用距今300年左右墓葬出土的云南现代人成年男性和女性头骨各30例作为研究材料,利用3D激光扫描技术及逆向工程软件提取头骨断面的外轮廓信息,在此基础上,采用几何形态测量方法对头骨冠状面、矢状面的外轮廓形状的性别差异进行了比较。研究结果显示:男女头骨在冠状面、正中矢状面顶骨部、正中矢状面枕骨部的形状上性别差异不显著,而在正中矢状面额骨部的形状上差异显著。本文对于古人类学和体质人类学鉴定人类头骨的性别差异,以及探讨不同地区、不同时代人群头面部形状的变异具有一定的参考价值。     

Exploring sexual dimorphism of human occipital and temporal bones through geometric morphometrics in an identified Western-European sample

Sex estimation is a paramount step of bioprofiling in both forensic anthropology and osteoarchaeology. When the pelvis is not optimally preserved, anthropologists commonly rely on the cranium to accurately estimate sex. Over the last decades, the geometric morphometric (GM) approach has been used to determine sexual dimorphism of the crania, in size and shape, overcoming some difficulties of traditional visual and metric methods. This article aims to investigate sexual dimorphism of the occipital and temporal region through GM analysis in a metapopulation of 50 Western-European identified individuals. Statistical analyses were performed to compare centroid size and shape data between sexes through the examination of distinct functional modules. Regression and Procrustes ANOVA were used to examine allometric and asymmetrical implications. Discriminant functions, combining size and shape data, were established. Significant dimorphism in size was found, with males having larger crania, confirming the major influence size has on cranial morphology. Allometric relationships were found to be statistically significant in both right and left temporal bones while shape differences between sexes were only significant on the right temporal bone. The visualization of the mean consensus demonstrated that males displayed a larger mastoid process associated with a reduced mastoid triangle and less projected occipital condyles. This exploratory study confirms that GM analysis represents an effective way to quantitatively capture shape of dimorphic structures, even on complex rounded ones such as the mastoid region. Further examination in a larger sample would be valuable to design objective visualization tools that can improve morphoscopic sex estimation methods.     

Sexual dimorphism in the skull of minks Mustela vison, badgers Meles meles and otters Lutra lutra

Multiple-group principal component analysis and discriminant analysis were used to investigate the morphological differences between adult skulls of male and female minks, badgers and otters from Norway. The first principal component axis, calculated from the variance-covariance matrix of log-transformed data, was interpreted as a growth-free size axis in all three species, while the other components were interpreted as representing shape. Having largely separated size and shape variation, these two aspects of sexual dimorphism could be studied. The standardized component scores were subjected to an analysis of variance and discriminant analyses were performed on size-in and size-out data. Sexual dimorphism was disclosed on eight of the 12 components in minks and on seven of the 12 components in badgers and otters. In mink the multivariate differences were more due to size than to shape, whereas in badgers and otters most of the multivariate differences were due to shape, but the differences in size were also significant. The shape dimorphism was shown to be functionally related to jaw and neck muscles. The results were discussed in relation to recent theories to explain the evolutionary significance of sexual dimorphism in body size of mustelids. It was concluded that these theories do not fully explain the dimorphism found in the skulls of the moderately dimorphic badger and otter.     

Changes in the sexual dimorphism of the human mandible during the last 1200 years in Central Europe

According to many investigations, changes in mandibular morphology can occur synchronously with changes in the environment, and sexual dimorphism of the mandible can be influenced by the environment. Sexual dimorphism during the last 1200 years was evaluated using geometric morphometric analysis of virtual cranial models. The method of geometric morphometrics allows differences in size and shape to be assessed separately. We analyzed groups of adult individuals dating to Early Middle Ages, High Middle Ages, Early Modern Ages and from a modern Czech population (21st century). Significant sexual dimorphism in mandibular size was found in all populations. A trend in the sexual dimorphism of size was seen, with differences between the sexes increasing gradually over time. Size changes in female mandibles were a better reflection of environmental conditions and climate than size changes in male mandibles. Regarding changes in the sexual dimorphism of shape, significant dimorphism was found in all four samples. However, the pattern of mandibular shape dimorphism was different and varied considerably between samples. There was only one stable shape trait showing sexual dimorphism across all four samples in our study: the gonion lies more laterally in male than in female mandibles and male mandibles are relatively wider than female mandibles. Sexual dimorphism of shape is not influenced by the climate; instead sexual selection might play a role. This research supports earlier studies that have found that the degree and pattern of sexual dimorphism is population-specific and the factors regulating sexual dimorphism today may not be the same as those in the past.     

Sexual dimorphism in Southeast Asian crania: A geometric morphometric approach

Despite a number of studies stating that sexual dimorphism is population specific, sexual differences in Southeast Asian populations have received little attention. Previous studies in this region have focused on samples from Thailand or East Asian populations from China and Japan, examining sexual dimorphism predominantly of the postcranial bones, teeth and mandible with comparatively few cranial studies. These earlier studies have used traditional methods to metrically assess differences between the sexes. The aim of this study is to use geometric morphometric methods for the first time to quantify sexual dimorphism of Southeast Asian crania and extend knowledge of cranial sexual dimorphism beyond China, Japan and Thailand. A total of 35 unilateral and midline landmark coordinates were collected from 144 mainland and island Southeast Asian crania (89 male, 55 female). Using the shape analysis software Morphologika, principal components analysis and thin plate splines allowed for the statistical and visual exploration of shape differences. Differences included relative facial breadth, particularly across the zygomatic and postorbital regions and cranial vault breadth. Significant size dimorphism was also apparent. Overall expected accuracies were highest in the discriminant analysis using both shape and centroid size (86.8%).     

Brief communication: Sexual dimorphism of the juvenile basicranium

The purpose of this article is to examine the level of sexual dimorphism exhibited in the foramen magnum and occipital condyles of juveniles, and to test the utility of this sexual dimorphism for estimating sex. Using five basicranial measurements taken from 36 juveniles of known sex and age from the Lisbon documented collection (Portugal), we evaluated sexual dimorphism in the juvenile cranial base. Our application of a method previously applied solely to adults indicated that the basicranium is sexually dimorphic in juveniles, with larger foramen magnum and occipital condyle dimensions observed in males. Significant univariate differences between males and females were found for length and breadth of the foramen magnum, and breadth of the left occipital condyle. Using these measurements, multivariate discriminant analysis indicated that sex was correctly assigned 75.8% of time. Obtained accuracy, however, was lower than reported by previous studies of adult samples. We suggest that this discrepancy is a result of population variability rather than age. Am J Phys Anthropol 2010. © 2009 Wiley‐Liss, Inc.     

Sexual dimorphism of Dall's porpoise and harbor porpoise skulls

To evaluate and quantify sexual dimorphism of skull shape and assess the ontogenetic background for differences, samples of 134 harbor porpoise (Phocoena phocoena) and 85 Dall's porpoise (Phocoenoides dalli) were compared in terms of cranial shape and shape ontogeny using three-dimensional geometric morphometrics. After correction for allometry, no sexual differences were detected in harbor porpoise, while Dall's porpoise showed statistically significant sexual dimorphism of skull shape. Since no sex-specific differences were detected in the directionalities of the ontogenetic vectors, we cannot reject that the dimorphism is innate. Based on the different mating systems of the two species and the lack of sexual dimorphism in the harbor porpoise, the dimorphism in Dall's porpoise is most likely a result of sexual selection in relation physical competition for mates given that male skulls provide room for larger neck muscles with a more favorable lever arm.     

Geometric morphometrics of the skull of Tinamidae (Aves, Palaeognathae)

The Tinamidae comprise exclusively Neotropical palaegnathous birds, with homogeneous body morphology and no sexual dimorphism. The goal of this work was to explore the variation in skull morphology between taxa and its possible correspondence with features such as diet or gender using geometric morphometric tools. Eleven landmarks were analyzed in 53 skulls of 4 genera that inhabit grasslands: Nothoprocta, Eudromia, Nothura and Rhynchotus. Intrageneric and intergeneric variability was analyzed. The genera studied here can be distinguished based on the geometric shape of their skull, with prenarial region length and neurocranium shape as the most outstanding features. In the genus Eudromia, males and females could be differentiated, while in the genus Nothoprocta, the species differentiated according to their trophic habits. This study allows establishing that genera and, in some cases, the gender of the Tinamidae can be differentiated based on cranial shape.     

实验室条件下唐鱼两性异形及其与游泳能力关系

为了解唐鱼两性异形及其与游泳能力关系,检测了性成熟阶段唐鱼躯干部和鱼鳍形态特征以及爆发游泳速度(U_burst)和临界游泳速度(U_crit)在雌雄之间的差异,旨在从形态适应角度探究长期进化中雌雄唐鱼各自面对选择压力所产生的游泳能力差异及其机制,从而为野生唐鱼保护提供基础数据.结果表明: 雌性唐鱼的体长、头高、头宽、尾鳍面积以及吻端至枕骨后末端、腹鳍起点至背鳍末端等长度均与雄性无显著差异.而体高、体宽、腹鳍起点至背鳍起点等反映腹腔大小的形态参数以及吻端至背鳍起点、吻端至臀鳍起点、枕骨后末端至背鳍起点等反映躯干部大小的形态参数均显示为雌性显著大于雄性,但头长以及胸鳍面积、腹鳍面积、背鳍面积和臀鳍面积均显示为雄性显著大于雌性.对所有数据进行主成分分析,结果显示第1主成分贡献率为74.2%,负载量较大的是体长、头长、头高、体高、头宽、体宽以及各鳍之间距离等主要反映唐鱼躯干整体特征的参数;第2主成分贡献率为15.7%,负载量较大的是胸鳍面积、腹鳍面积、背鳍面积和臀鳍面积等主要反映鱼鳍特征的参数.唐鱼性别在第1主成分上无法区分,但在第2主成分却可以明显区分.根据体宽、胸鳍面积、腹鳍面积、背鳍面积和臀鳍面积等建立的性别判别方程对雌雄判断准确率达到91.8%～92.5%.唐鱼游泳能力测定结果显示,雌性U_burst与雄性无显著差异,但U_crit显著小于雄性.以上结果表明,雌雄唐鱼两性异形主要集中在与游泳能力相关的鱼鳍特征上.相比雄性,雌性唐鱼虽然胸鳍等鱼鳍面积较小导致其U_crit小于雄性,却具有更长的躯干部以保证其同样具有较高的爆发游泳能力,从而有利于在流速波动很大的溪流中躲避捕食和进行其他应急活动;相比雌性,雄性唐鱼则具有较大的鱼鳍面积保证其U_crit高于雌性,以利于日常活动及在繁殖过程中追逐雌性等相对持久性游泳运动.     

Multivariate morphometries and sexual dimorphism in the orb-web spider Metellina segmentata (Clerck, 1757) (Araneae, Metidae)

Sexual dimorphism in body size and leg length was investigated in a common orb-weaving spider of Ireland and northern Europe, Metellina segmentata (Clerck, 1757) (Araneae, Metidae). Univariate and multivariate analyses of sexual dimorphism revealed that a greater proportion of between sex variation (sexual dimorphism) was attributable to variation in shape than in size. Significant differences were found in the scores for males and females for the first two principal components. PCI (shape) accounted for 44.25% of the variation and PC2 (size) 13.01% of the variation. Although M. segmentata has been attributed with minimal sexual size dimorphism, females were markedly heavier, possibly a reflection of differential reproductive investment between the sexes, but males had markedly longer legs and broader prosoma. The results are discussed with regard to existing theories of natural and sexual selection, particularly those concerning sexual cannibalism and differential life history traits in males and females. Models that attempt to explain the evolution of sexual size dimorphism in spiders and of the web builders in particular, fail to account for the multivariate nature of dimorphism, especially with respect to shape.     

Shape,size, and maturity trajectories of the human ilium

Morphological traits of the ilium have consistently been more successful for juvenile sex determination than have techniques applied to other skeletal elements, however relatively little is known about the ontogeny and maturation of size and shape dimorphism in the ilium. We use a geometric morphometric approach to quantitatively separate the ontogeny of size and shape of the ilium, and analyze interpopulation differences in the onset, rate and patterning of sexual dimorphism. We captured the shape of three traits for a total of 191 ilia from Lisbon (Portugal) and London (UK) samples of known age and sex (0–17 years). Our results indicate that a) there is a clear dissociation between the ontogeny of size and shape in males and females, b) the ontogeny of size and shape are each defined by non‐linear trajectories that differ between the sexes, c) there are interpopulation differences in ontogenetic shape trajectories, which point to population‐specific patterning in the attainment of sexual dimorphism, and d) the rate of shape maturation and size maturation is typically higher for females than males. Male and female shape differences in the ilium are brought about by trajectory divergence. Differences in size and shape maturation between the sexes suggest that maturity may confound our ability to discriminate between the sexes by introducing variation not accounted for in age‐based groupings. The accuracy of sex determination methods using the ilium may be improved by the use of different traits for particular age groups, to capture the ontogenetic development of shape in both sexes. Am J Phys Anthropol 156:19–34, 2015 © 2014 Wiley Periodicals, Inc.     

Ontogeny of facial dimorphism and patterns of individual development within one human population

Based on a longitudinal study of radiographs of the Denver Growth Study, we investigated the morphological development of individual and gender differences in the anterior neurocranium, face, and basicranium. In total, 500 X-rays of 14 males and 14 females, each with 18 landmarks and semilandmarks, were digitized and analyzed using geometric morphometric methods. Sexual dimorphism in shape and form is already present at the earliest age stage included in the analysis. However, the nature of dimorphism changes with age. Four factors apper to contribute to cranial sexual dimorphism in human postnatal development: 1) initial, possibly prenatal, differences in shape; 2) differences in the association of size and shape; 3) male hypermorphosis; and 4) some degree of difference in the direction of male and female growth trajectories. Studying changes in individuals, we find a low correlation between newborn and adult morphology, while 3-year-olds already show a high correlation with their adult form. We conclude that the adult pattern of interindividual difference in facial form in a single human population is established within the first few years of life.     

Morphometrical considerations on occipital condyles

From the study of the surface area of the occipital condyles and of skull volume resulted that the surface area is not proportional to skull volume. The measurements were conducted on a collection of 424 male and 317 female skulls. It was observed that the two condyles are rarely symmetrical and that also flat condyles appear. Bi-partitioned condyles can be divided into three types: roof-like bipartitions, simple bipartitions and total bipartitions.     

An ontogenetic perspective on the study of sexual dimorphism, phylogenetic variability, and allometry of the skull of European ground squirrel, Spermophilus citellus (Linnaeus, 1766)

Most studies of morphological variability in or among species are performed on adult specimens. However, it has been proven that knowledge of the patterns of size and shape changes and their covariation during ontogeny is of great value for the understanding of the processes that produce morphological variation. In this study, we investigated the patterns of sexual dimorphism, phylogenetic variability, and ontogenetic allometry in the Spermophilus citellus with geometric morphometrics applied to cross-sectional ontogenetic data of 189 skulls from three populations (originating from Burgenland, Banat, and Dojran) belonging to two phylogenetic lineages (the Northern and Southern). Our results indicate that sexual dimorphism in the ventral cranium of S. citellus is expressed only in skull size and becomes apparent just before or after the first hibernation because of accelerated growth in juvenile males. Sexes had the same pattern of ontogenetic allometry. Populations from Banat and Dojran, belonging to different phylogroups, were the most different in size but had the most similar adult skull shape. Phylogenetic relations among populations, therefore, did not reflect skull morphology, which is probably under a significant influence of ecological factors. Populations had parallel allometric trajectories, indicating that alterations in development probably occur prenatally. The species’ allometric relations during cranial growth showed characteristic nonlinear trajectories in the two northern populations, with accelerated shape changes in juveniles and continued but almost isometric growth in adults. The adult cranial shape was reached before sexual maturity of both sexes and adult size after sexual maturity. The majority of shape changes during growth are probably correlated with the shift from a liquid to a solid diet and to a lesser degree due to allometric scaling, which explained only 20 % of total shape variation. As expected, viscerocranial components grew with positive and neurocranial with negative allometry.     

Sex-linked variations in the sagittal otolith biometry of Nemipterus randalli (Russell, 1986) from the eastern Mediterranean Sea

Few studies have been conducted on the sagittal otolith shape and morphometry of Nemipterus randalli, and none of these studies has examined the effect of sexual dimorphism on the otolith morphology of this species, therefore this study aimed to contribute to knowledge about the otolith morphology of N. randalli, an invasive fish species for the Mediterranean Sea. For this purpose, a total of 132 samples (51 female and 81 male) were obtained from İskenderun Bay with the help of commercial fishermen in November 2018. Relationships between otolith measurements and fish size were determined. Shape indices and elliptic Fourier coefficients were calculated. Significant differences were detected between males and females in all analysis. The sexes were separated from each other using both shape indices and elliptic Fourier coefficients. However, shape analysis was more effective in distinguishing sexes than traditional morphometric analysis. Asymmetry in the otolith morphology of sexes has been attributed to differences in the growth and sexual maturity of male and female fish. The results of this study indicated that sexual dimorphism in Nemipterus randalli was also reflected in the otolith morphology.     

Sexual size and shape evolution in European newts (Amphibia: Caudata: Salamandridae) on the Balkan Peninsula

We used a phylogenetic perspective in an examination of the direction and extent of sexual dimorphism in body size and body shape in European newts from the Balkan Peninsula (alpine newts, Mesotriton alpestris; crested newts, Triturus cristatus superspecies; smooth newts, Lissotriton vulgaris). We found a strong, female‐biased sexual size dimorphism (SSD) in the analysed clades of alpine newt, whereas within crested newts we found a less stringent female‐biased SSD in Triturus carnifex, Triturus macedonicus and Triturus karelinii, and no significant SSD in T. cristatus or Triturus dobrogicus. Among the smooth newts, we found male‐biased SSD in Lissotriton vulgaris vularis and Lissotriton vulgaris greacus and no SSD in Lissotriton vulgaris meridionalis. Most of these newts also exhibit a significant sexual dimorphism in body shape, which varied more randomly than body size, regardless of SSD level. Female and male body size as well as the degree of SSD displayed statistically significant phylogenetic signal, while sexual dimorphism in body shape was phylogenetically independent. The relationship between independent contrast data for female size and male size indicated that SSD in European newts could be driven by a disproportionate increase in female size as increase in female size was not accompanied by a proportional increase in male size.     

Thin-plate spline analysis of allometry and sexual dimorphism in the human craniofacial complex

The relationship between allometry and sexual dimorphism in the human craniofacial complex was analyzed using geometric morphometric methods. Thin-plate splines (TPS) analysis has been applied to investigate the lateral profile of complete adult skulls of known sex. Twenty-nine three-dimensional (3D) craniofacial and mandibular landmark coordinates were recorded from a sample of 52 adult females and 52 adult males of known age and sex. No difference in the influence of size on shape was detected between sexes. Both size and sex had significant influences on shape. As expected, the influence of centroid size on shape (allometry) revealed a shift in the proportions of the neurocranium and the viscerocranium, with a marked allometric variation of the lower face. Adjusted for centroid size, males presented a relatively larger size of the nasopharyngeal space than females. A mean-male TPS transformation revealed a larger piriform aperture, achieved by an increase of the angulation of the nasal bones and a downward rotation of the anterior nasal floor. Male pharynx expansion was also reflected by larger choanae and a more posteriorly inclined basilar part of the occipital clivus. Male muscle attachment sites appeared more pronounced. In contrast, the mean-female TPS transformation was characterized by a relatively small nasal aperture. The occipital clivus inclined anteriorly, and muscle insertion areas became smoothed. Besides these variations, both maxillary and mandibular alveolar regions became prognathic. The sex-specific TPS deformation patterns are hypothesized to be associated with sexual differences in body composition and energetic requirements.     

Geographic variation in size and sexual dimorphism of North American weasels

Geographic variation in size (skull length) and sexual dimorphism in Mustela erminea, Mustela frenata and Mustela nivalis in North America is described and analysed in relation to latitude, longitude, climatic variables, and sympatry or allopatry of these species. Only erminea increases in size with latitude; it does so regardless of the presence or absence of frenata or nivalis. Latitude is a better predictor of size in erminea than available measures of climate, seasonality or prey size. There is no evidence for character displacement between any pair of species. The sexes covary in size in frenata and erminea , and probably in nivalis , although geographic variation in sexual dimorphism occurs in frenata and erminea. The principal cause of sexual dimorphism appears to be sexual selection for large size in males rather than the high energetic requirements resulting from an elongate body shape. However, prey size may constrain female size (and possibly also male size). Regional differences in the abundance of prey during the growth of young weasels may affect adult size much more in males than in females and contribute to geographic variation in sexual dimorphism.