Sexual dimorphism and cultural evolution in the Late Pleistocene and Holocene of Europe

Dental, cranial and body size data are reviewed for European Upper Paleolithic, Mesolithic and Neolithic males and females. Over these three periods there is a substantial decrease in the level of sexual dimorphism. From separate analysis of trends occurring between males and females, it is shown that the major cause for this decrease in sexual dimorphism is gracilization of the males between the Upper Paleolithic and Mesolithic. Reduction in males is related to shifting technological patterns associated with hunting and changes in the types of animals hunted. Further reduction in sexual dimorphism between the Mesolithic and Neolithic and from the Neolithic to modern European populations is shown to be more closely tied to changes occurring among females. Analysis of changing patterns of sexual dimorphism in Late Pleistocene and Holocene populations of Europe suggests an interrelationship between cultural and biological evolution.     

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.     

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.     

Sexual differences in Neanderthal limb bones

European and Near Eastern Neanderthal postcranial remains have been analyzed to determine the degrees of sexual dimorphism in limb bone size and robusticity present among the Neanderthals. The remains were sexed on the basis of pelvic morphology where possible (seven males and three females) and otherwise on the basis of absolute size employing limb bone lengths and articular dimensions (12 males and 15 females). Neanderthal sexual size dimorphism, both within single site samples and in the total sexable sample, is virtually the same as that of recent human samples. Furthermore, despite a tendency towards more robust limbs, the Neanderthals exhibit sexual dimorphism in limb bone shaft and articular robusticity similar to that of recent human samples. By the time of the Neanderthals, sexual dimorphism in limb bone size and robusticity appears to have reached recent human proportions.     

Skeletal indicators of subsistence patterns and activity régime in the Mesolithic sample from Grotta dell’Uzzo (Trapani,Sicily): a case study

The skeletal sample from Grotta dell’Uzzo, Sicily (2 adult females and 5 adult males) was compared to a number of more representative population samples from Western Europe and the Mediterranean Basin. The majority of these were from Italian pre- and protohistoric sites. The research protocol analyzed skeletal indicators of labour activity and sexual division of labour (body size and proportions, sexual dimorphism, limb lateralization, bone robustness, the development of muscular attachments, accessory articular facets, signs of muscular hyperfunction). Sexual dimorphism and limb lateralization showed some regular patterns of possible general significance in all the samples examined here. A general pattern of gracilization and de-specialization of physical activity is observed in the Mesolithic as compared to the Upper Palaeolithic samples. The main features of the Mesolithic samples are: a reduction of body size and bone robustness, a lower degree of sexual dimorphism and limb bone asymmetry, a reduction of the mechanical stress on the lower limbs indicated by less pronounced muscular attachments and reduced talar flattening. This trend is reversed towards the Neolithic period. The main features of these variations are discussed in relation to economic and environmental changes. The Uzzo sample fits well into the general picture of the Western European Mesolithic, although showing some intermediate features between the Mesolithic and the Neolithic samples. This paper is dedicated to the memory of the late Roland Menk, who made significant contribution to our understanding of the Mesolithic transition.     

Temporal changes in human temporomandibular joint size and shape

Measurements approximating the size of the temporomandibular joint were taken on a series of genetically homogeneous populations from early Nubia, which span almost 10,000 years and embody a shift from a primarily hunting and gathering adaptation to a completely agricultural lifeway. A generalized trend of reduction of temporomandibular joint size was observed. In addition, a decrease in sexual dimorphism was apparent for all measures of joint size; a change primarily mitigated by reductions in male dimensions. The observed variation in size and form of the temporomandibular joint is most likely the result of the reduction in masticatory muscle robusticity and resultant changes in craniofacial form which have been documented for the transition from a hunting and gathering to an agricultural subsistence in Nubia.     

Sexual dimorphism in postcranial skeletal shape suggests male-biased specialization for physical competition in anthropoid primates

Sexual dimorphism often arises as a response to selection on traits that improve a male's ability to physically compete for access to mates. In primates, sexual dimorphism in body mass and canine size is more common in species with intense male–male competition. However, in addition to these traits, other musculoskeletal adaptations may improve male fighting performance. Postcranial traits that increase strength, agility, and maneuverability may also be under selection. To test the hypothesis that males, as compared to females, are more specialized for physical competition in their postcranial anatomy, we compared sex-specific skeletal shape using a set of functional indices predicted to improve fighting performance. Across species, we found significant sexual dimorphism in a subset of these indices, indicating the presence of skeletal shape sexual dimorphism in our sample of anthropoid primates. Mean skeletal shape sexual dimorphism was positively correlated with sexual dimorphism in body size, an indicator of the intensity of male–male competition, even when controlling for both body mass and phylogenetic relatedness. These results suggest that selection on male fighting ability has played a role in the evolution of postcranial sexual dimorphism in primates.     

Sexual selection on skeletal shape in Carnivora

Lifetime reproductive success of males is often dependent upon the ability to physically compete for mates. However, species variation in social structure leads to differences in the relative importance of intraspecific aggression. Here, we present a large comparative dataset on sexual dimorphism in skeletal shape in Carnivora to test the hypotheses that carnivorans exhibit sexual dimorphism in skeletal anatomy that is reflective of greater specialization for physical aggression in males relative to females and that this dimorphism is associated with the intensity of sexual selection. We tested these hypotheses using a set of functional indices predicted to improve aggressive performance. Our results indicate that skeletal shape dimorphism is widespread within our sample. Functional traits thought to enhance aggressive performance are more pronounced in males. Phylogenetic model selection suggests that the evolution of this dimorphism is driven by sexual selection, with the best‐fitting model indicating greater dimorphism in polygynous versus nonpolygynous species. Skeletal shape dimorphism is correlated with body size dimorphism, a common indicator of the intensity of male–male competition, but not with mean body size. These results represent the first evidence of sexual dimorphism in the primary locomotor system of a large sample of mammals.     

Unusually low sexual dimorphism of endocranial capacity in a Zulu cranial series

The mean cranial capacities of 50 male and 50 female Zulu crania were found to be 1373.3 +/- 107.4 ml for males and 1251.2 +/- 101.1 ml for females (means +/- SD). The male value resembles that of other Negro groups, while the female value is somewhat higher than the value for Negro crania as a whole. The index of sexual dimorphism is 8.9%, which is low when compared with those of other Negroid series and other populations. The possible causes for this form of a low sexual dimorphism are as follows: A negative secular trend, with the assumption that the Zulu crania were larger than those of the reference populations of African Negroids before the start of the secular trend change. This would seem to be the most likely possibility, with some supporting evidence for both parts of the explanation. An absence of secular trend, with a demographic sampling aberration, in which large females and small males of the population are sampled. This possibility cannot be totally excluded. An absence of secular trend, with a genetic difference in sexual dimorphism for cranial capacity between the Zulu and the reference populations. While this possibility cannot be excluded, it would be the least preferable explanation.     

Infanticide and fertility among Eskimos: A computer simulation

Until recently, certain Eskimo groups were reported to practice female infanticide in the belief that the time spent suckling a girl would delay the mother's next opportunity to bear a son, males being preferred to females because of their future role as providers in a hunting economy. From sex ratios in census data, rates of female infanticide of up to 66% for some groups have been inferred, leading some ethnographers to conclude that these groups were headed for extinction. Eskimo beliefs regarding the effects of infanticide on fertility, however, are in accord with the results of research on the relation of fertility and lactation: The cessation of lactation following infanticide would significantly shortern the expected interval until the next birth. Given this fact and available field data regarding the parameters of Eskimo population growth, the present computer simulation indicates that Eskimo populations could sustain a rate of 30% female infanticide and still survive. Higher reported rates are explained as the combined result of female infanticide plus the tendency of ethnographers to overestimate to overestimate the ages of juvenile females relative to juvenile males.     

Upper pleistocene fossil hominids from Sri Lanka

Between 1978 and 1983 hominid skeletal remains were collected from the cave sites of Batadomba lena and Beli lena Kitulgala in Sri Lanka (formerly Ceylon). These are the most ancient specimens of anatomically modern Homo sapiens found thus far in South Asia, radiocarbon dates placing them in the Upper Pleistocene. Morphometric analysis of the remains of some 38 individuals from the two sites indicates that these populations were characterized by medium stature, moderate to pronounced cranial and postcranial robusticity, medium-size permanent tooth crown measurements, prognathic alveolar facial proportions, and low incidence of osseous and dental pathological conditions. Comparisons of these ancient Sri Lankans with other prehistoric skeletal series from South Asia and elsewhere support the hypothesis that muscular-skeletal robusticity was a significant physical adaptation of earlier hunting-foraging populations. A trend towards reduction of sexual dimorphism and development of more gracile body form and smaller teeth appears to have accelerated with the socioeconomic transition to food-production strategies involving agriculture and pastoralism and refinement of technologies for food procurement and preparation, as documented by morphometric studies of later prehistoric inhabitants of South Asia.     

Morphological variation of introduced species: The case of American mink (Neovison vison) in Spain

We studied the morphology of American mink Neovison vison in five out of the six introduced populations in Spain. The spatial and temporal variation of body weight (BW), body length (BL), tail length, hind-foot length and ear length were analysed. Temporal trends in BW and BL in relation to years since mink introduction were also analyzed. In addition, we tested the effect of sex, age (juvenile, subadult and adult) and age–sex interaction, on each parameter. Morphological parameters differed between populations, illustrating the high variability of body size of American mink in different environments, and the phenotypic plasticity of the species. Annual variations were synchronized between populations, suggesting a large-scale effect on all of them. BW and BL showed a decreasing trend in both males and females in relation to years since introduction. This decrease may be related to mink's diet. Differences in sex and age were found, pointing to sexual dimorphism in adults, subadults and juveniles. The dimorphism in non-adult individuals suggests that subadult males may have a competitive advantage from subadult females in feeding and/or hunting on bigger prey from an early age (resource partitioning hypothesis).     

Sexual dimorphism in northern temperate spiders: implications for the differential mortality model

We examined sexual size dimorphism in 627 species from 123 genera and 32 families of northern temperate spiders from Great Britain and Ireland with different life histories, using phylogenetically independent contrasts. Web-building and non-web-building, sit-and-wait predators were compared with non-web-building, active hunting spiders. After accounting for phylogenetic effects, we find no evidence of differences in sexual size dimorphism in northern temperate spiders with differing life history/predatory strategies. We discuss the implications of our findings for the generality of the differential mortality hypothesis (Vollrath & Parker, 1992) with respect to spiders with different predatory modes from different habitats and environments. This recent theory proposed that extreme sexual dimorphism in spiders resulted from differential adult mortality as a consequence of different adult life histories. We conclude that this model cannot explain the less extreme dimorphism found in temperate spiders.     

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.     

Statistical study of sexual dimorphism in the human fetal sciatic notch

Whether human fetal skeletal remains exhibit sexual dimorphism has been the subject of considerable debate. Most attention in this debate has focused on the greater sciatic notch of the ilium, since it is a gross morphological characteristic with known sex differences in the adult and is easily seen in fetal skeletal remains. Unfortunately, previous traditional morphometric analyses of the fetal sciatic notch have led to ambiguous results. The purpose of this study is to determine whether differences between the sexes can be discerned when modern morphometric techniques are applied to the fetal sciatic notch. Photographs of the ventral side of 133 fetal ilia of known age and sex from the Trotter Collection of Washington University were digitized, and the trace coordinates used for all subsequent analyses. The results of the analysis demonstrate that there is significant sexual dimorphism in the anterior to posterior location of the maximum depth of the sciatic notch, but that the depth of the notch itself is not dimorphic. While there is significant sexual dimorphism in the shape of the sciatic notch, the amount of overlap between males and females is too great for the sciatic notch to be used as a reliable indicator of sex. © 1995 Wiley-Liss, Inc.     

Interspecific allometry for sexual shape dimorphism: Macroevolution of multivariate sexual phenotypes with application to Rensch's rule

Allometric trends in the degree of sexual dimorphism with body size have long fascinated evolutionary biologists. Many male-biased clades display more prominent sexual dimorphism in larger taxa (Rensch's rule), with most examples documenting this pattern for body size dimorphism. Although sexual dimorphism in traits other than body size is equally functionally relevant, characterizing allometric patterns of sexual dimorphism in such traits is hampered by lack of an analytical framework that can accommodate multivariate phenotypes. In this article, we derive a multivariate equivalency for investigating trends in sexual dimorphism—relative to overall body size—across taxa and provide a generalized test to determine whether such allometric patterns correspond with Rensch's rule. For univariate linear traits such as body size, our approach yields equivalent results to those from standard procedures, but our test is also capable of detecting trends in multivariate datasets such as shape. Computer simulations reveal that the method displays appropriate statistical properties, and an empirical example in Mediterranean lizards provides the first demonstration of Rensch's rule in a multivariate phenotype (head shape). Our generalized procedure substantially extends the analytical toolkit for investigating macroevolutionary patterns of sexual dimorphism and seeking a better understanding of the processes that underlie them.     

Relative growth,ontogeny, and sexual dimorphism in gorilla(Gorilla gorilla gorilla and G. g. beringei): Evolutionary and ecological considerations

Gorillas are the largest and among the most sexually dimorphic of all extant primates. While gorillas have been incorporated in broad-level comparisons among large-bodied hominoids or in studies of the African apes, comparisons between gorilla subspecies have been rare. During the past decade, however, behavioral, morphological, and molecular data from a number of studies have indicated that the western lowland (Gorilla gorilla gorilla) and eastern mountain (Gorilla gorilla beringei) subspecies differ to a greater extent than has been previously believed. In this study I compare patterns of relative growth of the postcranial skeleton to evaluate whether differences between subspecies result from the differential extension of common patterns of relative growth. In addition, patterns of ontogeny and sexual dimorphism are also examined. Linear skeletal dimensions and skeletal weight were obtained for ontogenetic series of male and female G.g. gorilla (n = 315) and G.g. beringei (n = 38). Bivariate and multivariate methods of analysis were used to test for differences in patterns of relative growth, ontogeny, and sexual dimorphism between sexes of each subspecies and in same-sex comparisons between subspecies. Results indicate males and females of both subspecies are ontogenetically scaled for postcranial proportions and that females undergo an earlier skeletal growth spurt compared to males. However, results also indicate that the onset of the female growth spurt occurs at different dental stages in lowland and mountain gorillas and that mountain gorillas may be characterized by higher rates of growth. Finally, data demonstrate lowland and mountain gorilla females do not differ significantly in adult body size, but mountain gorilla males are significantly larger than lowland gorilla males, suggesting mountain gorillas are characterized by a higher degree of sexual dimorphism in body size. Thus, although lowland and mountain gorillas do not appear to have evolved novel adaptations of the postcranium which correlate with differences in locomotor behavior, the present investigation establishes subspecies differences in ontogeny and sexual dimorphism which may be linked with ecological variation. Specifically, these findings are evaluated in the context of risk aversion models which predict higher growth rates and increased levels of sexual dimorphism in extreme folivores. Am. J. Primatol. 43:1–31, 1997. © 1997 Wiley-Liss, Inc.     

Sex ratio variation in harvested moose (<Emphasis Type="Italic">Alces alces</Emphasis>) calves: does it reflect population calf sex ratio or selective hunting?

During the last 30 years, the proportion of males in the calf harvest of moose (Alces alces) in Norway has decreased, indicating a decline in proportions of males recruited to the autumn populations. At the same time, the percentages of exclusive calf hunting permits and of calves shot have increased. The change in calf sex ratio may thus simply be the result of hunter preferences for slightly larger (6.2% higher body mass) male calves combined with fewer opportunities for selective hunting due to increasing hunting quotas of calves. We examined this hypothesis by analyzing the variation in sex, number of siblings, carcass mass, date, and location of kill of 16,330 moose calves harvested during 1970–2004. In the presence of hunting selection for larger calves, we predicted larger proportions of male calves to be harvested in populations with large sexual size dimorphism among calves. Similarly, we expected more males to be harvested from twin than single litters because hunters then can more easily compare twins and select the larger calf, which is more often a male. Increasing proportions of single female calves were also expected to occur in the daily harvest as the accumulated number of harvested calves increased and the proportion of calves left in the population decreased. We found no positive relationship between the proportion of male calves and the level of sexual size dimorphism, no clear difference in sex ratio between harvested single and twin calves, and no increase in the proportion of single female calves as the accumulated number of calves in the harvest increased. This suggests that the spatiotemporal variation in the harvest calf sex ratio in Norway most likely reflects differences in population calf sex ratios prior to the hunting season and not varying degrees of hunting selectivity.     

Degree of pelvic sexual dimorphism in human populations. A greene t-test application

The knowledge of sexual dimorphisms is important in many aspects of paleodemography and paleobiology. One major problem lies in a correct sexual diagnosis. However, due to the different methodologies employed to estimate the degree of sexual dimorphism, the comparison of the results remains difficult. The Greene t-test (1989) is accurate for a comparative analysis of raw data. On the basis of 18 measurements of theos coxae the test is made to evaluate the degree of sexual dimorphism. The data collected are employed to investigate the pelvic sexual dimorphism within a recent sample of known sex, and among the Afalou-bou Rhummel — Taforalt (Epipaleolithic of Northern Africa) and the Teviec — Hoedic samples (Mesolithic of France). A comparative analysis is applied to a few recent samples. The results indicate the stability of sexual dimorphism pattern in recent and past populations.     

Relationships between lower limb cross-sectional geometry and mobility: the case of a Neolithic sample from Italy

This study investigates the relationships between lower limb robusticity and mobility in a Neolithic sample (LIG) from Italy (6th millennium BP). This study tests the hypothesis that the high femoral robusticity previously observed in the LIG sample is a consequence of the subsistence strategy (i.e., high mobility on uneven terrain) practiced by LIG. Cross-sectional geometric properties of the femur and tibia at midshaft of LIG (eight males and eight females) were collected and results compared to Late Upper Paleolithic (12 males, five females), Mesolithic (24 males, 8 females), and Eneolithic (28 males, 17 females) samples from other sites throughout Europe. The results show that the LIG sample does not show the reduction of lower limb robusticity that is characteristic of the Eneolithic sample, but rather that the LIG sample is most similar to the earlier, highly mobile, populations. This high level of robusticity in the LIG sample could reflect both their pastoral subsistence strategy combined with a rugged environment, as well as their earlier temporal position within the Neolithic. The results of this study further point to significant variation in male-female mobility patterns in the region, also possibly related to pastoral behavioral patterns.