Body Size, Weapon Use, and Natural Selection in the European Upper Paleolithic and Mesolithic

Evidence for a relationship between hunting strategies and body size is examined for human skeletons dating to the European Upper Paleolithic and Mesolithic. Trends for reduced limb size and stature seem to be correlated with improvements in the types of weapons utilized and a shift from aggressive to more docile game. Although some of these observations fit the predictions of Brues concerning the spearman-archer model, it is suggested that selection for reduced metabolic demands is a more plausible explanation for decrease in body size from the Upper Paleolithic to the Mesolithic . [Upper Paleolithic, Mesolithic, hunting, body size]     

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.     

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.     

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.     

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.     

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.     

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.     

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.     

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).     

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.     

Is human longevity a consequence of cultural change or modern biology?

Increased longevity, expressed as the number of individuals surviving to older adulthood, represents a key way that Upper Paleolithic Europeans differ from earlier European (Neandertal) populations. Here, we address whether longevity increased as a result of cultural/adaptive change in Upper Paleolithic Europe, or whether it was introduced to Europe as a part of modern human biology. We compare the ratio of older to younger adults (OY ratio) in an early modern human sample associated with the Middle Paleolithic from Western Asia with OY ratios of European Upper Paleolithic moderns and penecontemporary Neandertals from the same region. We also compare these Neandertals to European Neandertals. The difference between the OY ratios of modern humans of the Middle and Upper Paleolithic is large and significant, but there is no significant difference between the Neandertals and early modern humans of Western Asia. Longevity for the West Asian Neandertals is significantly more common than for the European Neandertals. We conclude that the increase in adult survivorship associated with the Upper Paleolithic is not a biological attribute of modern humans, but reflects important cultural adaptations promoting the demographic and material representations of modernity.     

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.     

Le Paléolithique supérieur au Japon

The Japanese archipelago has been colonized by humans in different ways and at different times: it has therefore undergone a succession of varied continental influences. However, the origin of these populations would be mainly northern and located at the beginning of the Upper Paleolithic, in the European conception. Subsequently, at least two other continental migratory waves were observed, through the center and then through the south of the archipelago. This latter influence brought the “Mesolithic” population still active today in the northern islands of Hokkaido.     

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.     

Regression equations for estimating stature from long bones of early Holocene European samples

Regression equations for estimating living stature from long bone lengths have been calibrated on a sample of European Neolithic skeletons (33 males and 27 females) by using both least-squares (model I) and major-axis (model II) regression techniques. Stature estimates of the skeletal sample have been made by means of Fully's anatomical method, a procedure based on the sum of all osseous components of height, providing the best approximations to the actual stature. The calculated equations have been tested, along with those generally used to predict stature of earlier European remains, on a small, well-preserved sample including Late Upper Paleolithic, Mesolithic, and Neolithic skeletons. The results indicate that the model II equations are particularly useful when very short or very tall individuals are involved and, at the same time, are among the best predictors of stature in less extreme conditions. © 1996 Wiley-Liss, Inc.     

欧洲旧石器时代晚期石器类型学评介及类型学相关问题探讨

类型学是考古材料描述与分析的基础。近年来,学者们对类型学尤其是旧石器时代石器类型学的反思较多,涉及类型学的理论基础、分类原则、类型学的考古解释能力等各个方面,然而少有中国学者系统介绍西方常用的旧石器考古类型学体系。鉴于此,本文主要介绍D. de Sonneville-Bordes与J. Perrot建立且在欧洲、西亚及北非应用较为普遍的旧石器时代晚期石器类型学,并思考旧石器考古学中类型学应用的相关问题。由于文化的区域多样性,描述中国的旧石器考古学材料时显然不能照搬欧洲的类型学体系,但可参考它并建立适宜于中国旧石器时代考古材料描述的体系。若此,一方面,我们可以更好地将中国的旧石器晚期考古材料与其他地区的相关材料进行对比;另一方面,可以揭示中国旧石器时代晚期石器类型的区域多样化。     

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.     

Microevolution of outer and inner structures of upper molars in Late Pleistocene and Early Holocene humans

In this study, we investigate outer and inner variations of upper second molars (UM2) for Late Pleistocene and Early Holocene modern humans, at a key-period in our evolutionary history associated with major sociocultural, economic and environmental changes. Non-metric traits have been recorded on 89 UM2 of 66 Upper Paleolithic, Mesolithic and Neolithic individuals, and 40 UM2 have been microscanned to assess variations in enamel thickness (ET) distribution and enamel–dentine junction (EDJ) shape. Major changes are found between Mesolithic and Neolithic periods: a decrease of the metacone expression combined with an increase of the hypocone development; an increase of the heterogeneity of ET distribution between lingual and buccal cusps; and an increase of the development of the dentine horn tips corresponding to the hypocone and, to a lesser extent, to the metacone. These morphological modifications could be linked to the masticatory functional changes associated with the transition to agriculture.     

Evolution of the supraorbital region in Upper Pleistocene fossil hominids from South-Central Europe

South-Central European fossil hominids dated to the Upper Pleistocene exhibit a distinct morphological and metric continuum in supraorbital form from early Neandertal (Krapina), through late Neandertals (Vindija), to early Upper Paleolithic hominids. The supraorbital morphologies pertinent to this continuum are documented, and the alterations in size and morphology are discussed ralative to the function of supraorbital superstructures and their relationship to overall craniofacial form. It is concluded that this continuum most likely reflects localized transition between Neandertals and modern man in this region of Europe.     

Radiocarbon dating: The continuing revolution

Radiocarbon (¹⁴C) dating, now in its fifth decade of general use, continues to be the most widely employed method of inferring chronometric age for late Pleistocene and Holocene age materials recovered from archeological contexts. Over the last decade, several technical advances in ¹⁴C studies have provided contexts for a number of significant applications in archeology that were previously either not possible or not practical. These include the extension of the calibrated ¹⁴C time scale into the late Pleistocene and the development of accelerator mass spectrometry (AMS). The contribution of AMS-based ¹⁴C values to the critical evaluation of archeological data is illustrated by considering the problems of dating early plant domestication in the Near East and Mesoamerica, New World Paleoindian human skeletal materials, and European Upper Paleolithic and Mesolithic materials.