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.     

Robusticity and sexual dimorphism in the postcranium of modern hunter-gatherers from Australia

Throughout much of prehistory, humans practiced a hunting and gathering subsistence strategy. Elevated postcranial robusticity and sexually dimorphic mobility patterns are presumed consequences of this strategy, in which males are attributed greater robusticity and mobility than females. Much of the basis for these trends originates from populations where skeletal correlates of activity patterns are known (e.g., cross-sectional geometric properties of long bones), but in which activity patterns are inferred using evidence such as archaeological records (e.g., Pleistocene Europe). Australian hunter-gatherers provide an opportunity to critically assess these ideas since ethnographic documentation of their activity patterns is available. We address the following questions: do skeletal indicators of Australian hunter-gatherers express elevated postcranial robusticity and sexually dimorphic mobility relative to populations from similar latitudes, and do ethnographic accounts support these findings. Using computed tomography, cross-sectional images were obtained from 149 skeletal elements including humeri, radii, ulnae, femora, and tibiae. Cross-sectional geometric properties were calculated from image data and standardized for body size. Australian hunter-gatherers often have reduced robusticity at femoral and humeral midshafts relative to forager (Khoi-San), agricultural/industrialized (Zulu), and industrialized (African American) groups. Australian hunter-gatherers display more sexual dimorphism in upper limb robusticity than lower limb robusticity. Attributing specific behavioral causes to upper limb sexual dimorphism is premature, although ethnographic accounts support sex-specific differences in tool use. Virtually absent sexual dimorphism in lower limb robusticity is consistent with ethnographic accounts of equivalently high mobility among females and males. Thus, elevated postcranial robusticity and sexually dimorphic mobility do not always characterize hunter-gatherers.     

Head shape allometry and proximate causes of head sexual dimorphism in Podarcis lizards: joining linear and geometric morphometrics

Podarcis bocagei and Podarcis carbonelli are two lacertid species endemic to the western Iberian Peninsula, and both show head size and shape sexual dimorphism. We studied immature and adult head sexual dimorphism and analysed ontogenetic trajectories of head traits with body and head size, aiming to shed light on the proximate mechanisms involved. Immatures were much less dimorphic than adults, but geometric morphometric techniques revealed that head shape sexual differences are already present at this stage. Males and females differed in allometry of all head characters with body size, with males showing a disproportionate increase of head size and dimensions. On the other hand, head dimensions and head shape changed with increasing head size following similar trends in both sexes, possibly indicating developmental restrictions. Consequently, adult sexual dimorphism for head characters in these species is the result of both shape differences in the immature stage and hypermetric growth of the head in relation to body size in males.  © 2008 The Linnean Society of London, Biological Journal of the Linnean Society , 2008, 93 , 111–124.     

Labrador Inuit Subsistence in the Context of Environmental Change: An Initial Landscape History Perspective

In this article, I examine the subsistence economy of the contact-period Labrador Inuit (17th to 19th century) to investigate the relationships of elements of specific cultural changes to environmental change, the effects of cultural interactions, and internal social processes. I apply paleoclimate and sea-ice records and zooarchaeological studies of life history and seasonality of ecologically sensitive species to reconstruct histories of change in particular elements of the physical landscape and of their use. I associate the use of communal houses by the Labrador Inuit and their modification of settlement patterns during the 18th century to a limited phase of environmental moderation and stability during the Little Ice Age and suggest that these changes reflect, in part, economic strategies oriented toward surplus production.     

Sexual variation in assimilation efficiency: its link to phenotype and potential role in sexual dimorphism

Sex-specific variation in morphology (sexual dimorphism) is a prevalent phenomenon among animals, and both dietary intake and resource allocation strategies influence sexually dimorphic traits (e.g., body size or composition). However, we investigated whether assimilation efficiency (AE), an intermediate step between dietary intake and allocation, can also vary between the sexes. Specifically, we tested whether sex-based differences in AE can explain variation in phenotypic traits. We measured morphometric characteristics (i.e., body length, mass, condition, and musculature) and AE of total energy, crude protein, and crude fat in post-reproductive adult Children’s pythons (which exhibit a limited female-biased sexual size dimorphism) fed both low and high dietary intakes. Meal size was negatively related to AE of energy. Notably, male snakes absorbed crude protein more efficiently and increased epaxial (dorsal) musculature faster than females, which demonstrates a link between AE and phenotype. However, females grew in body length faster but did not absorb any nutrient more efficiently than males. Although our results do not provide a direct link between AE and sexual size dimorphism, they demonstrate that sexual variation in nutrient absorption exists and can contribute to other types of sex-based differences in phenotype (i.e., sexual dimorphism in growth of musculature). Hence, testing the broader applicability of AE’s role in sexually dimorphic traits among other species is warranted.     

Sexual dimorphism and human tooth size differences

Current male/female differences in tooth size are due to the male/female differences in body bulk that exist in any given human population. These differences are residues of the sexual dimorphism that was maintained for adaptive reasons during the Middle Pleistocene. Late in the Pleistocene the development of food processing techniques led to the reduction of both male and female dental dimensions. Dental sexual dimorphism, however, was maintained until the very end of the Pleistocene when the hunting of large game animals by crude techniques was replaced by a focus on great numbers of small game caught by more sophisticated means and by an increasing utilization of plant foods. The subsequent reduction in dimorphism represents the actions of the Probable Mutation Effect operating under conditions of relaxed selection. The conclusion offered is that the smallest degree of sexual dimorphism visible in the modern world is to be found among those populations that are separated by the greatest interval of time from precursors who depended for their survival on a Pleistocene big game hunting mode of subsistence.     

Variation in guenon skulls (II): sexual dimorphism

Patterns of size and shape sexual dimorphism in adult guenons were examined using a large sample of skulls from almost all living species. Within species, sexual dimorphism in skull shape follows the direction of size-related shape variation of adults, is proportional to differences in size, and tends to be larger in large-bodied species. Interspecific divergence among shape trajectories, which explain within species sex differences, are small (i.e., trajectories of most species are nearly parallel). Thus, changes in relative proportions of skull regions that account for the distinctive shape of females and males are relatively conserved across species, and their magnitude largely depends on differences in size between sexes. A conservative pattern of size-related sexual dimorphism and a model of interspecific divergence in shape which strongly reflects size differences suggest a major role of size and size-related shape variation in the guenon radiation. It is possible that in the guenons, as in the neotropical primates (with whom they have obvious parallels), size has helped to determine morphological change along lines of least evolutionary resistance, influencing sexual dimorphism. In Miopithecus and Erythrocebus, the smallest and largest guenon genera, it is likely that the interaction of ecology and size contributes significantly to patterns of sexual dimorphism. The results of this study thus emphasise the need to consider allometry and size alongside ecology and behaviour when examining primate sexual dimorphism.     

Development of intraspecific size variation in black coucals,white-browed coucals and ruffs from hatching to fledging

Most studies on sexual size dimorphism address proximate and functional questions related to adults, but sexual size dimorphism usually develops during ontogeny and developmental trajectories of sexual size dimorphism are poorly understood. We studied three bird species with variation in adult sexual size dimorphism: black coucals (females 69% heavier than males), white-browed coucals (females 13% heavier than males) and ruffs (males 70% heavier than females). Using a flexible Bayesian generalized additive model framework (GAMM), we examined when and how sexual size dimorphism developed in body mass, tarsus length and bill length from hatching until fledging. In ruffs, we additionally examined the development of intrasexual size variation among three morphs (Independents, Satellites and Faeders), which creates another level of variation in adult size of males and females. We found that 27–100% of the adult inter- and intrasexual size variation developed until fledging although none of the species completed growth during the observational period. In general, the larger sex/morph grew more quickly and reached its maximal absolute growth rate later than the smaller sex/morph. However, when the daily increase in body mass was modelled as a proportion, growth patterns were synchronized between and within sexes. Growth broadly followed sigmoidal asymptotic models, however only with the flexible GAMM approach, residual distributions were homogeneous over the entire observation periods. These results provide a platform for future studies to relate variation in growth to selective pressures and proximate mechanisms in these three species, and they highlight the advantage of using a flexible model approach for examining growth variation during ontogeny.     

GROWTH AND SEXUAL DIMORPHISM OF ATLANTIC WALRUSES (ODOBENUS ROSMARUS ROSMARUS) IN FOXE BASIN, NORTHWEST TERRITORIES, CANADA

Growth of Atlantic walruses ( Odobenus rosmarus rosmarus ) was investigated using morphological data collected in association with Inuit subsistence walrus hunts. Four growth models were examined. The growth parameters of a constrained Richards model were used to quantify growth and to test for sexual dimorphism. The asymptotic length of male walruses (315.2 cm ± 3.8 (SE), n = 103) was significantly larger ( t = 7.21, df = 191, P < 0.05) than the asymptotic length of females (276.6 cm ± 3.4, n = 90). Sexual size dimorphism in adults was due to a longer growth period and a faster growth rate in males. The predictive equation relating mass ( M , kg) to standard length ( SL , cm) was: Log₁₀ M = -3.74 + 2.68(Log₁₀ SL ), n = 25, r ²= 0.98. There were no significant differences in the size of male walruses from Foxe Basin collected in the 1950s and this study. There were too few data to compare females. There were no significant differences in size between walruses sampled in Greenland and Foxe Basin in the 1980s and 1990s. Foxe Basin walruses were significantly larger than walruses sampled in northern Hudson Bay in the 1950s. Female Atlantic walruses sampled in Foxe Basin were larger than female Pacific walruses ( Odobenus rosmarus divergens ) sampled in Alaska.     

Is sexual dimorphism in the femur a "population specific phenomenon"?

The patterns of sexual dimorphism as well as the differences in amount between the populations were studied on a sample of 162 male and 159 female left femora, which were classified as Zulu, Sotho, Xosa and South Africans of European extraction. Multivariate analyses revealed that even adjacent African tribes exhibit a different pattern of sexual dimorphism, but there were similarities between Zulu and European femora. Furthermore, relative size differences, i.e. shape, discriminated more clearly between the sexes than did absolute size. Bicondylar width yielded a statistically significant higher degree of sexual dimorphism in Europeans when compared to African populations. This finding was interpreted in terms of the biomechanical demands on the femur under different living conditions. On the other hand, sexual dimorphism of femoral length did not differ among the populations. This was unexpected since femoral length correlates highly with stature, which was reported to show a lesser degree of sexual dimorphism in Africans than in Europeans. Detailed analyses of the results of the present study led to suggest that different living conditions may affect bones in complex ways of which linear growth is only one aspect.     

Sexual dimorphism in human lower limb bone structure: relationship to subsistence strategy and sexual division of labor

Cross-sectional geometric properties of the human femur and tibia are compared in males and females in a number of recent and archaeological population samples extending back to the Middle Paleolithic. There is a consistent decline in sexual dimorphism from hunting-gathering to agricultural to industrial subsistence strategy levels in properties which measure relative anteroposterior bending strength of the femur and tibia in the region about the knee. This trend parallels and is indicative of reductions in the sexual division of labor, in particular differences in the relative mobility of males and females. Sexual dimorphism in mediolateral bending strength near the hip shows no consistent temporal trend, probably reflecting relatively constant sex differences in pelvic structure related to the requirements of childbirth. Upper and Middle Paleolithic samples are indistinguishable in terms of sexual dimorphism from modern huntergatherers, suggesting a similar sexual division of labor. The results illustrate the utility of cross-sectional geometric parameters of long bone diaphyses in reconstructing behavioral differences within and between past populations. Some variations in the accuracy of sexing techniques based on diaphyseal measurements of the lower limb long bones may also be explained by these behavioral and structural factors.     

Sex-limited mutations and the evolution of sexual dimorphism

Abstract.— Although the developmental and genetic mechanisms underlying sex differences are being elucidated in great detail in a number of species, there remains a breach between proximate and evolutionary studies of sexual dimorphism. More precisely, the evolution of sex-limited gene expression at autosomal loci has not been well reasoned using either theoretical or empirical methods. Here, I show that a Mendelian genetic model including elementary details of sexual differentiation provides novel insight into the evolution of sex differences via sex limitation. This model indicates that the nature of allelic effects and the pattern of selection must be known in both sexes to predict the evolution of sex differences. That is, selection interacts with genetic variation for sexual dimorphism to produce unanticipated patterns of trait divergence or convergence between the sexes. Ultimately, this model may explain why previous models for the evolution of sexual dimorphism do not predict the erratic behavior of the sex difference during artificial selection experiments.     

Effect of mobility on femur midshaft external shape and robusticity

This study investigates differences in femur midshaft shape, robusticity, and sexual dimorphism derived from external measurements between a broad range of prehistoric and historic North American populations with different subsistence strategies and inferred levels of mobility. The sample was divided into six groups to test whether observed femur midshaft variables follow the patterns predicted based on archaeologically and historically determined subsistence and mobility data. The results suggest significant variation in femur midshaft shape and robusticity in all populations, and that inferred mobility levels do not correspond consistently with femur midshaft structure in either males or females. Results do, however, support the prediction that sexual dimorphism is generally greater in more mobile populations.     

Patterns of sexual dimorphism in the hominoid distal humerus

Basic biomechanical principles predict that body size differences and differences in the positional behavior of primates should impact on the design of the locomotor skeleton. Allometric distortions in joint shape might be expected between sexes if the degree of body size dimorphism is substantial and/or if sex-specific differences exist in behavior. Nevertheless, there are few documented cases of sexual dimorphism in the limb joints of hominoids, despite substantial body size dimorphism and some reports of intersexual differences in positional behavior. This study re-examines sexual dimorphism in the hominoid distal humerus using coordinate data, and distinguishes explicitly between degree of dimorphism (i.e., the magnitude of intersexual differences) and pattern of dimorphism (i.e. , the nature of these differences). Using a variety of multivariate morphometric methods (e.g., canonical variates analysis of Mosimann shape variables; Euclidean Distance Matrix Analysis of both form and pattern difference matrices), we address the following issues: (1) do males and females of different species and subspecies (or ethnic groups for humans) maintain similar joint shapes? (2) are multiple patterns of dimorphism evident in this region of hominoids? (3) are differences and similarities in degree and pattern predicted by phylogenetic propinquity and positional behavior? For the most part, our results support earlier findings that sexual dimorphism in the shape of the anthropoid elbow is slight. Of the eight taxa considered here, only the western lowland gorillas exhibited significant differences in the shape of the distal humerus. Gorilla gorilla gorilla also displays a significantly different pattern of dimorphism from the orang-utan. Pattern differences between Andaman Islanders and both mountain gorillas and the orang-utan also approach statistical significance (P<0.06 and P<0.08, respectively). Overall, and despite marked differences in the degree of dimorphism, the knuckle-walking African apes are more similar in patterns of dimorphism to each other than to other taxa (e.g., gorillas are more similar to orang-utans in degree, but more similar to chimpanzees and bonobos in pattern). We could find no definitive "human pattern" in our results and suspect that this is because human upper limbs face less stringent mechanical constraints since they are relieved of locomotor stresses (but we cannot rule out the possibility of undocumented differences among our human groups in sex-specific, work-related activities). We anticipate finding additional pattern differences among anthropoids in articular dimorphism as we add other taxa to our sample (including fossil hominids), and examine other joint systems.     

Sexual differences in larval life history traits of acanthocephalan cystacanths

Sexual differences in life history traits, such as size dimorphism, presumably arise via sexual selection and are most readily observed in adults. For complex life-cycle parasites, however, sexual selection may also have consequences for larval traits, e.g., growth in intermediate hosts. Two acanthocephalan species (Acanthocephalus lucii and Echinorhynchus borealis) were studied to determine, whether larval life histories differ between males and females. The size of female A. lucii cystacanths had a much stronger relationship with intermediate host size than males, suggesting females invest more in growth and are consequently more limited by resources. No relationship between host size and cystacanth size was observed for E. borealis. For both species, female cystacanths survived longer in a culture medium composed entirely of salts, which could suggest that females have greater energy reserves than males. A comparative analysis across acanthocephalan species indicated that sexual size dimorphism at the adult stage correlates with cystacanth dimorphism. However, the relationship was not isometric; cystacanths do not reach the same level of sexual dimorphism as adults, possibly due to resource constraints. Our results suggest that larval life histories diverge between males and females in some acanthocephalans, and this is seemingly a consequence of sexual selection acting on adults.     

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.     

Incorporating an ontogenetic perspective into evolutionary theory of sexual size dimorphism

Sexual size dimorphism (SSD) describes divergent body sizes of adult males and females. While SSD has traditionally been explained by sexual and fecundity selection, recent advances in physiology and developmental biology emphasize that SSD would occur proximately because of sexual differences in ontogenetic growth trajectories (i.e., growth rate and duration). Notably, these ontogenetic traits are subject to energetic or time constraints and thus traded off with fitness components (e.g., survival and reproduction). To elucidate the importance of such ontogenetic trade‐offs in the evolution of SSD, we developed a new theoretical framework by extending quantitative genetic models for the evolution of sexual dimorphism in which we reinterpret the trait as body size and reformulate sex‐specific fitness in size‐dependent manners. More specifically, we assume that higher growth rate or longer growth duration leads to larger body size and higher reproductive success but incurs the cost of lower survivorship or shorter reproduction period. We illustrate how two sexes would optimize ontogenetic growth trajectories in sex‐specific ways and exhibit divergent body sizes. The present framework provides new insights into the evolutionary theory of SSD and predictions for empirical testing.     

Which proximate factor determines sexual size dimorphism in tiger snakes?

Diverse interactions between factors that influence body size complicate the identification of the primary determinants of sexual size dimorphism. Using data from a long‐term field study (1997–2009), we examined the contributions of the main proximate factors potentially influencing sexual size dimorphism from birth to adulthood in tiger snakes (Notechis scutatus). Data on body size, body mass and body condition of neonates, juveniles and adults were obtained by mark–recapture. Frequent recaptures allowed us to monitor reproductive status, diet and food intake, and to estimate survival and growth rates in age and sex classes. Additional data from females held briefly in captivity enabled us to assess reproductive output and the body mass lost at parturition (proxies for reproductive effort). From birth to maturity, individuals of both sexes experienced similar growth and mortality rates. We found no difference in diet, feeding and survival rates between the sexes, nor between juveniles and adults. On maturity, despite comparable diet and food intake by both sexes, the high energy requirements of vitellogenesis and gestation were responsible for a depletion of body reserves and probably resulted in a marked decrease in growth rates. Males were largely exempt from such costs of reproduction, and so could grow faster than females and attain larger body sizes. The absence of niche divergence between the sexes (uniformity of habitat, lack of predators) suggests that the impact of differential energetic investment for reproduction on growth rate is probably the main proximate factor influencing sexual size dimorphism in this species. © 2011 The Linnean Society of London, Biological Journal of the Linnean Society, 2011, 103 , 668–680.     

Differential sexual dimorphism: size and shape in the cranium and pelvis of grey foxes (Urocyon)

Patterns of sexual size dimorphism (SSD) and cranial dimorphism are well documented. However, limited examinations exist of the contrasts in the patterns and nature of dimorphism across body regions (e.g. cranium, pelvis), particularly when these regions have different sex-specific functions (e.g. display in mating, locomotion, and reproduction). Using landmark-based morphometric techniques, we investigated size and shape dimorphism variation in the crania and pelves of two closely-related fox species within the genus Urocyon . Although we found no significant size and shape dimorphism in the crania of either species, we did find significant dimorphism in the pelvis: its size was dimorphic in Urocyon littoralis (but not in Urocyon cinereoargenteus ) and its shape was dimorphic in both species (though more pronounced in U. littoralis ). The observation of greater dimorphism in the pelvis than in the cranium suggests that factors such as offspring size and locomotor mode play a greater role in sexual dimorphism than simple 'whole body' allometric affects associated with dimorphism in body size.  © 2009 The Linnean Society of London, Biological Journal of the Linnean Society , 2009, 96 , 339–353.     

Bioanthropological analysis of human occipital condyles using geometric morphometric method

Sex differences are present in all parts of the body, including the skeletal system. Several methods are used to analyze the sex differences of skeleton, while more recently, a new method called geometric morphometry has been used. The aim of this study was to examine the sexual dimorphism of occipital condyles on human skulls originating from the population of Bosnia and Herzegovina using the geometric morphometric method.Material and methodsThe study was conducted on 214 human skulls of known gender from Bosnian population. For analysis of sexual dimorphism of occipital condyles, we used geometric morphometry, where all the skulls were scanned to obtain three-dimensional skull models. On the obtained models, we marked anthropometric points on occipital condyles in a Landmark Editor program from which we exported data in the form NTSYS file and analyzed it in MorphoJ program.ResultsFirst principal component PC1 describes 26.917% of total variability, the second principal component PC2 describes 20.992% of total variability, while the first eight principal components together describe 100% of total variability. The greatest variability between the male skulls and female skulls was present in the anterior-posterior diameter (length of occipital condyles). Discriminant functional analysis of the shape and size of the occipital condyles was possible with 69.50% accuracy for male skulls and with 60.27% accuracy for female skulls. The size of the occipital condyles showed a statistically significant effect on sexual determination. Discriminant functional analysis of the shape of the occipital condyles without affecting size enabled the determination of gender with with 65.96% accuracy for male skulls and with 63.01% accuracy for female skulls.ConclusionAnalysis of sexual dimorphism of occipital condyles using geometric morphometry showed statistically significant differences in the shape and size of occipital condyles between the sexes. The accuracy of sex determination based on occipital condyles was higher for male gender.