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.     

Metric dental change in the European upper paleolithic and mesolithic.

Evolutionary trends for dental reduction are presented for European Upper Paleolithic and Mesolithic samples. The analysis demonstrates that the greatest decrease in tooth size occurs between the two divisions of the Upper Paleolithic, while little and insignificant change characterizes the Late Upper Paleolithic/Mesolithic transition. Trends for tooth size over this period indicate that (1) human evolution does not stop with the appearance of "anatomically modern Homo sapiens," (2) changes in tooth size fluctuate with increases in the efficiency and complexity of cultural systems, and (3) the Early Upper Paleolithic sample should be considered transitional between Wurm II European Neanderthals and later Upper Paleolithic and Mesolithic groups.     

Post-pleistocene reductions in human dental structure: a reappraisal in terms of increasing population density

This paper discusses the well-documented acceleration which occurred in the reduction of human dental structure during the post-Pleistocene. It suggests that the process can be explained in terms of different but related factors inherent in a transition from late-Pleistocene hunting and gathering conditions to ones involving a sedentary life in larger groups. It is postulated that directional selective forces acting during Upper Paleolithic to maintain large tooth size had only a poor influence and that the new techniques in food preparation had little or no effect. The phenomenon is seen as a possible side-effect of a more complex overall reduction in body size, of which dental reduction only represents a small but demonstrable part. It is also suggested that stresses may have played an important role in producing a transitory reduced dimensional expression of the genetic background. An interaction between the variables directly or indirectly affecting body mass is shown in the model proposed which relates to post-Pleistocene. An increase in population density is indicated as being the most decisive biological factor in determining the acceleration in the trend towards reduction. A preliminary version of this paper was presented at the Symposium on Upper Paleolithic/Mesolithic of Europe and the Mediterranean Basin, Pisa, September 8–10, 1984.     

Sexual dimorphism in relation to big-game hunting and economy in modern human populations

Postcranial skeletal data from two recent Eskimo populations are used to test David Frayer's model of sexual dimorphism reduction in Europe between the Upper Paleolithic and Mesolithic. Frayer argued that a change from big-game hunting and adoption of new technology in the Mesolithic reduced selection for large body size in males and led to a reduction in skeletal sexual dimorphism. Though aspects of Frayer's work have been criticized in the literature, the association of big-game hunting and high sexual dimorphism is untested. This study employs univariate and multivariate analysis to test that association by examining sexual dimorphism of cranial and postcranial bones of two recent Alaskan Eskimo populations, one being big-game (whale and other large marine mammal) hunting people, and the second being salmon fishing, riverine people. While big-game hunting influences skeletal robusticity, it cannot be said to lead to greater sexual dimorphism generally. The two populations had different relative sexual dimorphism levels for different parts of the body. Notably, the big-game hunting (whaling) Eskimos had the lower multivariate dimorphism in the humerus, which could be expected to be the structure under greatest exertion by such hunting in males. While the exertions of the whale hunting economic activities led to high skeletal robusticity, as predicted by Frayer's model, this was true of the females as well as the males, resulting in low sexual dimorphism in some features. Females are half the sexual dimorphism equation, and they cannot be seen as constants in any model of economic behavior. © 1993 Wiley-Liss, Inc.     

Body size, body proportions, and mobility in the Tyrolean "Iceman"

Body mass and structural properties of the femoral and tibial midshafts of the "Iceman," a late Neolithic (5,200 BP) mummy found in the Tyrolean Alps, are determined from computed tomographic scans of his body, and compared with those of a sample of 139 males spanning the European early Upper Paleolithic through the Bronze Age. Two methods, based on femoral head breadth and estimated stature/bi-iliac (pelvic) breath, yield identical body-mass estimates of 61 kg for the Iceman. In combination with his estimated stature of 158 cm, this indicates a short but relatively wide or stocky body compared to our total sample. His femur is about average in strength compared to our late Neolithic (Eneolithic) males, but his tibia is well above average. His femur also shows adaptations for his relatively broad body (mediolateral strengthening), while his tibia shows adaptations for high mobility over rough terrain (anteroposterior strengthening). In many respects, his tibia more closely resembles those of European Mesolithic rather than Neolithic males, which may reflect a more mobile lifestyle than was characteristic of most Neolithic males, perhaps related to a pastoral subsistence strategy. There are indications that mobility in general declined between the European Mesolithic and late Neolithic, and that body size and shape may have become more variable throughout the continent following the Upper Paleolithic.     

Mobility in Upper Paleolithic and Mesolithic Europe: evidence from the lower limb

A growing body of archeological evidence suggests that the dramatic climatic events of the Last Glacial Maximum in Europe triggered important changes in foraging behavior, involving a significant decrease in mobility. In general, changes in mobility alter patterns of bending of the midshaft femur and tibia, resulting in changes in diaphyseal robusticity and shape. This relationship between levels of mobility and lower limb diaphyseal structure was used to test the hypothesized decrease in mobility. Cross-sectional geometric data were obtained for 81 Upper Paleolithic and Mesolithic European femora and tibiae. The sample was divided into three time periods: Early Upper Paleolithic (EUP), Late Upper Paleolithic (LUP), and Mesolithic (Meso). In addition, because decreased mobility often results in changes in sex roles, males and females were analyzed separately. All indicators of bending strength decrease steadily through time, although few of the changes reach statistical significance. There is, however, a highly significant change in midshaft femur shape, with LUP and Meso groups more circular in cross-section than the EUP sample, supporting archeologically based predictions of decreased mobility. Sexual dimorphism levels in diaphyseal strength remain low throughout the three time periods, suggesting a departure in Upper Paleolithic and Mesolithic foragers away from the pattern of division of labor by sex observed in modern hunter-gatherers. Results confirm that the onset of the Last Glacial Maximum represents a crucial stage in Late Pleistocene human evolution, and signals the appearance of some of the behavioral adaptations that are usually associated with the Neolithic, such as sedentism.     

Evolutionary trends of stature in upper Paleolithic and Mesolithic Europe

Long bone lengths of all available European Upper Paleolithic (41 males, 25 females) and Mesolithic (171 males, 118 females) remains have been transformed into stature estimates by means of new regression equations derived from Early Holocene skeletal samples using "Fully's anatomical stature" and the major axis regression technique (Formicola & Franceschi, 1996). Statistical analysis of the data, with reference both to time and space parameters, indicates that: (1) Early Upper Paleolithic samples (pre-Glacial Maximum) are very tall; (2) Late Upper Paleolithic groups (post-Glacial Maximum) from Western Europe, compared to their ancestors, show a marked decrease in height; (3) a further, although not significant, reduction of stature affects Western Mesolithics; (4) no regional differences have been observed during both phases of the Upper Paleolithic; (5) a high level of homogeneity has also been found in the Mesolithic, both in Western and Eastern Europe; (6) the internal homogeneity found during the Mesolithic in Western and Eastern Europe is associated with marked inter-regional variability, with populations of the latter region showing systematically significantly greater stature than their Western contemporaries. Evaluation of possible causes for the great stature of the Early Upper Paleolithic samples points to high nutritional standards as the most important factor. Results obtained on later groups clearly indicate that the Last Glacial Maximum, rather than the Mesolithic transition, is the critical phase in the negative trend affecting Western European populations. While changes in the quality of the diet, and in particular decreased protein intake, provide a likely explanation for that trend, variations in levels of gene flow probably also played a role. Reasons for the West-East Mesolithic dichotomy remain unclear and lack of information for the Late Upper Paleolithic of Eastern Europe prevents insight into the remote origins of this phenomenon. Analysis of regional differentiation of stature, particularly well supported by data from Mesolithic sites, points to the absence of today's latitudinal gradients and suggests a relative homogeneity in dietary, cultural and biodemographic patterns for the last hunter-gatherer populations of Western Europe.     

Subsistence activities and the sexual division of labor in the European Upper Paleolithic and Mesolithic: Evidence from upper limb enthesopathies

Studies of cultural artifacts and faunal remains from European Upper Paleolithic and Mesolithic sites indicate a shift in hunter gatherer subsistence strategies, involving an intensification and diversification of resource exploitation relative to earlier foragers during the Tardiglacial and Postglacial periods. This trend has been recognized as well through the analysis of non-pathological skeletal adaptations of the upper limbs of European Upper Paleolithic human fossils. These paleoanthropological studies of adaptive bone modeling also raise the question of female use of throwing-based weapon technology in the Upper Paleolithic. Here, we studied another type of osteological marker of activity, enthesopathies, of the upper limb remains of 37 European Upper Paleolithic and Mesolithic human fossils, with the goal of testing two hypotheses: 1) that activity levels were heightened at the end of Upper Paleolithic and into the Mesolithic relative to earlier foragers of the Gravettian, and 2) that there was an absence of a marked sexual division of labor in European hunter-gatherers during this time span. Our results are consistent with the first hypothesis; upper limb enthesopathies are significantly less frequent in the Gravettian group, but raise doubts about the second hypothesis. Four males exhibit lesions that can be confidently associated with throwing activities, while no females exhibit such lesions.     

On the evolution of diet and landscape during the Upper Paleolithic through Mesolithic at Franchthi Cave (Peloponnese, Greece)

Franchthi Cave in southern Greece preserves one of the most remarkable records of socioeconomic change of the Late Pleistocene through early Holocene. Located on the southern end of the Argolid Peninsula, the area around the site was greatly affected by climate variation and marine transgression. This study examines the complex interplay of site formation processes (material deposition rates), climate-driven landscape change, and human hunting systems during the Upper Paleolithic through Mesolithic at Franchthi Cave based on the H1B faunal series. Building on earlier work, we establish the full spectrum of the meat diet using taphonomic evidence, and we analyze these data for trends in socioeconomic reorganization. Foraging patterns during the Aurignacian and “Gravettoid” occupations at Franchthi were terrestrial and already rather diversified in comparison to Middle Paleolithic diets in southern Greece. Hunting shifted abruptly to a mixed marine-terrestrial pattern during the Final Paleolithic, and fishing activities intensified though the Mesolithic. The zooarchaeological data indicate two consecutive trends of increasing dietary breadth, the first within an exclusively terrestrial context, and the second as marine habitats came into use through the end of the Mesolithic. The intensity of the human occupations at this site increased in tandem with intensified use of animal and plants. Comparison to the inland site of Klissoura Cave 1 indicates that the trend toward broader diets was regional as well as local.     

Animal resources and subsistence strategies

Several faunal assemblages excavated in deposits of different antiquity (from Lower Paleolithic to Bronze Age), located in Northern, Central and Southern Italy, were studied from the archeozoological and taphonomic point of view. Data obtained by different Authors allow reconstruction of subsistence strategies adopted by prehistoric humans in these areas and through time, in particular as far as the exploitation of animal resources is concerned. The following assemblages were considered: Isernia La Pineta (Molise; Lower Paleolithic), Grotta Breuil (Latium; Middle Paleolithic), Grotta della Ghiacciaia (Verona; Middle Paleolithic), Riparo di Fumane and Riparo Tagliente (Verona; Middle and Upper Paleolithic), Riparo Mochi (Liguria; Upper Paleolithic), Grotta della Continenza (L'Aquila; Upper Paleolithic and Mesolithic), Grotta dell'Edera (Trieste; Mesolithic and Neolithic), Grotta della Cala at Marina di Camerota (Salerno; Eneolithic), Contraguda (Sassari; Neolithic), Castellaro Lagusello (Mantova; Bronze Age). Exploitation of the vegetal resources has been analyzed in the Neolithic sites of Colle Santo Stefano (Fucino), Settefonti (L'Aquila) and Catignano (Pescara).     

Hearth-side socioeconomics, hunting and paleoecology during the late Lower Paleolithic at Qesem Cave, Israel

The late Lower Paleolithic archaeofaunas of Qesem Cave in the southern Levant span 400-200 ka and associate with Acheulo-Yabrudian (mainly Amudian) industries. The large mammals are exclusively Eurasian in origin and formed under relatively cool, moist conditions. The zooarchaeological findings testify to large game hunting, hearth-centered carcass processing and meat sharing during the late Lower Paleolithic, not unlike the patterns known from Middle and Upper Paleolithic caves in the region. Well-defined hearth features are rarely preserved in Qesem Cave, but the heterogeneous distributions of burned bones indicate areas of frequent hearth rebuilding throughout the occupation sequence. The hominins delayed consumption of high quality body parts until they could be moved to the cave, where hearths were hubs of processing activities and social interaction. Paradoxically, the cut marks on the Qesem bones are both more abundant and more randomly oriented than those observed in Middle and Upper Paleolithic cases in the Levant. These results suggest that several individuals were directly involved in cutting meat from the bones and that the social mechanics of meat sharing during the late Lower Paleolithic at Qesem Cave differed from those typical of both the Middle and Upper Paleolithic in the region.     

中国旧石器时代石球的实验研究

石球是一种旧石器时代分布极为广泛的石质工具,有关其形制、制作与使用还存在较大的探索空间。其形制包括一般打制石球与琢制滚圆的石球。中国发现石球按大小可以分为两个类型,较大者多属于旧石器时代早中期,较小者多属于旧石器时代晚期及以后时期,本文讨论侧重前者。实验研究显示,石球生产是一项时间成本高昂的活动,如果加上琢制过程,所需要时间远多于手斧生产。旧石器时代早中期石球的大小与质量存在明显的偏向性,与成年男性手掌大小一致,这支持石球可能是直接握持使用的。投掷使用实验表明,平均质量与大小、形制规整的石球最有利于用手抓握投掷。结合考古材料中石球集中发现的状况与伴生动物化石进一步支持石球很可能是一种狩猎工具。不过采用石球狩猎距离较短,杀伤力有限,且更依赖机会,有较高风险。     

Brachial and crural indices of European late Upper Paleolithic and Mesolithic humans

Among recent humans brachial and crural indices are positively correlated with mean annual temperature, such that high indices are found in tropical groups. However, despite inhabiting glacial Europe, the Upper Paleolithic Europeans possessed high indices, prompting Trinkaus (1981) to argue for gene flow from warmer regions associated with modern human emergence in Europe. In contrast, Frayer et al. (1993) point out that Late Upper Paleolithic and Mesolithic Europeans should not exhibit tropically-adapted limb proportions, since, even assuming replacement, their ancestors had experienced cold stress in glacial Europe for at least 12 millennia. This study investigates three questions tied to the brachial and crural indices among Late Pleistocene and recent humans. First, which limb segments (either proximal or distal) are primarily responsible for variation in brachial and crural indices? Second, are these indices reflective of overall limb elongation? And finally, do the Late Upper Paleolithic and Mesolithic Europeans retain relatively and/or absolutely long limbs? Results indicate that in the lower limb, the distal limb segment contributes most of the variability to intralimb proportions, while in the upper limb the proximal and distal limb segments appear to be equally variable. Additionally, brachial and crural indices do not appear to be a good measure of overall limb length, and thus, while the Late Upper Paleolithic and Mesolithic humans have significantly higher (i.e., tropically-adapted) brachial and crural indices than do recent Europeans, they also have shorter (i.e., cold-adapted) limbs. The somewhat paradoxical retention of "tropical" indices in the context of more "cold-adapted" limb length is best explained as evidence for Replacement in the European Late Pleistocene, followed by gradual cold adaptation in glacial Europe.     

Inorganic raw materials economy and provenance of chipped industry in some stone age sites of northern and central Italy

An opportunistic and local choice of raw materials is typically attested in the Lower and Middle Paleolithic industries throughout Italy. The quality of the raw material usually affected the flaking technology and quality of the products. In the Upper Paleolithic and the Mesolithic, raw material procurement strategies were more complex. Flint was exploited both locally, in areas where abundant outcrops of raw materials were available (such as the Lessini mountains), and in distant localities, after which it was transported or exchanged over medium/long distances. Different routes of exchange were thus followed in the various periods; good reconstruction of these routes have been provided by a study of the Garfagnana sites in Northern Tuscany, and the Mesolithic deposit of Mondeval de Sora (Dolomites). An interesting example of a Late Upper Paleolithic flint quarry and workshop were found in Abruzzo, in the San Bartolomeo shelter. The extended trade of obsidian from Lipari, Palmarola and Sardinia to the Italian Peninsula is attested in the Neolithic, with some differences concerning the age and different areas.     

Epipaleolithic human appendicular remains from Ein Gev I,Israel

The Upper Paleolithic (Early Epipaleolithic/Kebaran; ∼ 19,000 cal BP) human skeleton, from Layer 3 of Ein Gev I on the western flanks of the Golan Heights adjacent to the Sea of Galilee, retains sufficient limb remains to permit assessment of its body size and proportions, as well as diaphyseal reflections of skeletal hypertrophy. The individual was of modest stature but average mass for a later Upper Paleolithic individual, providing it with the body mass-to-stature body proportions characteristic of later Upper Paleolithic and more recent circum-Mediterranean humans. The humeri exhibit unexceptional diaphyseal asymmetry and robustness for an Upper Paleolithic human, and the femur exhibits similar relative diaphyseal hypertrophy. The humeral midshafts are relatively round, but the femoral and tibial midshafts are pronounced anteroposteriorly. As such, Ein Gev 1 provides additional paleobiological data on the appendicular remains of these Southwest Asian humans prior to the increasing sedentism of the terminal Pleistocene.     

Yugoslav Mesolithic dental reduction

Yugoslav Mesolithic dentition exhibits maximum mesiodistal reduction compared with contemporary European and North African groups. This reduction is not explained entirely by attrition, and may be seen as a continuation of the European Upper Paleolithic trend. Buccolingual dimension does not reduce as much. In fact, this dimension in premolars and molars is larger than in other groups. This observation also occurs in Natufians, who were grain collectors, hunters, and gatherers. The Yugoslav Mesolithic group was collecting and domesticating Cerelia as well as fishing and hunting. Linear enamel hypoplasias (LEH) indicate childhood stress through the fifth year, which corroborates previously reported incidence of rickets in this group. The central maxillary incisors and canines manifest higher degrees of LEH, but the appearance on the second molars suggest a more severe physiological disruption. Sex differences in distributions of alveolar resorption and calculus suggest differences in diet or nutritional stress. Previous reports indicate that females had higher incidence of osteomalacia. If so, female nutritional stress may explain the extreme mesiodistal reduction and minimal sexual dimorphism in this group.     

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.     

The supra-acetabular fossa and groove: a skeletal marker for Northwest European mesolithic populations

A nonmetric trait, known as the supra-acetabular fossa and groove, is described for European Upper Paleolithic and mesolithic populations. This morphological feature consists of a pit posterior or superior to the anterior-inferior iliac spine and an associated groove which arcs over the roof of the acetabulum. Presence of the trait seems to be primarily related to the origin of the reflected head of rectus femoris. The trait is variable in its expression, shows an increase in definition with age, and occurs equally in males and females. Although the trait is commonly found in pre-Upper Paleolithic hominids, it is extremely rare in the Upper Paleolithic, limited to a single occurrence in the Oberkassel 2 female. In the Mesolithic, except for two innominates from Vlasac (Yugoslavia), the trait is found only in NW European populations, especially Skateholm (Sweden) and Henriksholm-Bøgebakken (Denmark) where it occurs in more than 50% of the known pelves. Given the trait’s restricted geographic distribution and high occurrence in Skateholm and Henriksholm-Bøgebakken, it is suggested the supra-acetabular fossa and groove represents a skeletal marker for NW European Mesolithic populations. As such, it provides evidence for the development of social territories in the early Holocene.     

Changes in the ‘Connectedness’ and Resilience of Paleolithic Societies in Mediterranean Ecosystems

Human predator‐prey relationships changed dramatically in the Mediterranean Basin between 250,000 and 9,000 years ago. Many of these changes can be linked to increases in Paleolithic human population densities. Small game species are particularly diagnostic of increases in human hunting pressure and are a major source of evidence for demographic change after 40–45,000 years ago. Biomass-corrected data on prey choice also indicate increasing use of those species that possess higher reproductive efficiencies. Step-wise, apparently irreversible shifts in human predatory niche are apparent in the Mediterranean Basin, beginning with the earliest Upper Paleolithic in the east and spreading westward. Evidence of demographic pressure and greater use of resilient prey populations is followed by technological innovations to exploit these animals more efficiently. The zooarchaeological findings suggest that Middle and Lower Paleolithic reproductive units probably were not robust at the micropopulation scale, due to the rather narrow set of behavioral responses that characterized social groups at the time, and thus localized extinctions at the micropopulation level were likely to have been common. Upper Paleolithic groups were the quintessential colonizers and, in addition, uniquely good at holding on to habitat gained. Upper Paleolithic archaeological “cultures” had shorter histories of existence than those of earlier periods, but they were even more widespread geographically. The demographic robustness of the Upper Paleolithic systems may stem from wholesale strategies for evening-out or sharing risk and volatility in technology. Micropopulations were larger and often denser on landscapes, more connected via cooperative ties, and thus more robust.     

The Azilian man from Los Azules Cave I (Cangas de Onis,Oviedo, Spain)

The author analyzes in this study the human skeleton found in the Azilian Grave of Los Azules Cave I (Asturias, NW of Spain), dated by C-14 between 7480 and 7590 B. C. It was diagnosed as an adult male, having a general strong robustness and high stature, as well as a congenital fusion of the right talus bones and, probably, bilateral claw foot. The comparison to the last Upper Paleolithic, the Epipaleolithic and the Mesolithic Human remains from Western Europe showed the similarity of Los Azules I Man to the robust Cromagnoids from these periods.