Regional variation in the postcranial robusticity of late Upper Paleolithic humans

Early modern humans from the European Upper Paleolithic (UP) demonstrate trends in postcranial biomechanical features that coincide with the last glacial maximum (LGM). These features have been interpreted as evidence that ecological changes of the LGM played a critical role in cultural and biological adaptation in European UP populations. In areas outside of Europe, similar environmental changes occurred with the LGM. This analysis introduces postcranial material from the Late Upper Paleolithic (LUP) of North Africa and Southeast Asia and tests two related hypotheses: 1) LUP samples across the Old World had similar patterns of postcranial robusticity and 2) relative to an available Early Upper Paleolithic (EUP) sample, regional LUP samples demonstrate similar trends in robusticity that may be attributable to climatic effects of the LGM. Cross-sectional geometric data of the humeri and femora were obtained for 26 EUP and 100 LUP humans from Europe, Africa, and Asia. Despite regional differences, LUP samples are similar relative to the EUP sample. In the humerus, bilateral asymmetry decreases in all LUP samples relative to the EUP sample. In the femur, LUP samples demonstrate increasingly circular femoral midshaft sections, reflecting reduced anteroposterior bending strength relative to the EUP sample. These patterns suggest changes in subsistence behavior and mobility after the LGM across the Old World that are most consistent with reduced mobility and broad-spectrum resource exploitation.     

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.     

Body size and postcranial robusticity of European Upper Paleolithic hominins

The robust diaphyses of Pleistocene hominins are said to indicate higher activity levels in these prehistoric humans than among people today. Thus, it could be argued that the prediction of body mass from fossil lower limb diaphyseal cortical area (CA) using recent human regressions might lead to erroneously high body mass estimates. This study uses three body mass prediction formulae based on the following features: reconstructed femoral 80% (subtrochanteric) CA, femoral head diameter (FH), and bi-iliac breadth and stature (BIB-St) among European Early and Late Upper Paleolithic (EUP and LUP) and recent humans from Africa and Europe. All three methods produce similar body mass estimates for all groups studied, including recent humans.Gleaning behavioral differences from these data is more difficult, as no significant differences in CA were found among the fossil and recent Europeans. It has been suggested that the EUP had less robust diaphyses than their LUP counterparts. However, here this result is only obtained when CA is size-standardized to femoral length(3) (Ruff et al., 1993, Am. J. phys. Anthrop.91, 21-53 Trinkaus et al., 1998, in Neandertals and Modern Humans in Western Asia, pp.391-404, New York: Plenum). This should not be interpreted as evidence for lower activity levels in the EUP, but rather as an artefact of standardization, for as Wolpoff (1999), Am. J. phys. Anthrop.109, 416-423 points out, these standardized variables are extremely sensitive to limb length differences, and the EUP have longer limbs than their LUP counterparts. With this in mind, these data do not support a pattern of behavioral differences between EUP and LUP humans, and therefore more sensitive measures than CA may be required to detect such differences.     

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.     

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.     

Interprétation paléogéographique des données palynologiques du site du Paléolithique supérieur de Mezhyrich (Ukraine)

The results of palynological study of the Upper Paleolithic Mezhyrich site in central Ukraine are presented. Four local pollen zones were identified. The results of the palynological investigations allow us to hypothesize that the plant cover during Upper Paleolithic had a mosaic structure. The pollen record shows the presence of a few tree species with the predominance of herbaceous plants. We suggest that the river valleys and other local favorable sites were natural refugia where forest elements could survive even during the Last Glacial Maximum. These favorable landscapes to the existence of prehistoric hunters may have persisted during the Last Pleniglacial.     

Mobility in Central European Late Eneolithic and Early Bronze Age: femoral cross-sectional geometry

Some scholars explain the absence of settlements in the Bohemian and Moravian Late Eneolithic (Corded Ware archaeological culture) as a consequence of pastoral subsistence with a high degree of mobility. However, recent archaeological studies argued that the archaeological record of the Late Eneolithic in Central Europe exhibits evidence for sedentary subsistence with mixed agriculture, similar to the subsequent Early Bronze Age. Because the archaeological data do not allow us to address unambiguously the mobility pattern in these periods, we used cross-sectional analysis of the femoral midshaft to test mobility directly on the human skeletal record. The results of femoral midshaft geometry do not support a high degree of mobility in the Late Eneolithic in Central Europe. This conclusion is supported mainly by no significant differences in male groups between the Late Eneolithic and Early Bronze Age in mechanical robusticity and shape of the femoral midshaft, although Corded Ware males still exhibit the highest absolute mean values of the diaphyseal shape (I(A-P)/I(M-L)) ratio and antero-posterior second moment of area. However, Late Eneolithic females have significantly higher torsional and overall bending rigidity because of a significantly higher medio-lateral second moment of area. This finding cannot be directly linked with a higher degree of long-distance mobility for these females. A significant difference was also found in overall decrease of size parameters of the femoral midshaft cross section for one of the Early Bronze Age samples, the Wieselburger females. Since the decrease of size and mechanical robusticity for Wieselburger females does not correspond with the parameters of Early Bronze Age females, we can expect a mosaic pattern of changes during the Late Eneolithic and Early Bronze Age period, instead of a simple unidirectional (diachronic) change of the mechanical environment.     

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.     

L'hominidé du Pléistocène supérieur en Corée

Since the middle of 1970s, one rock-shelter, six caves and one open archaeological site have yielded several hominid fossils. Among them, fossils from Yonggok, Mandal, Sangsi and Hungsu cave sites are useful to reconstruct the general shape of the Upper Pleistocene Hominids in Korea. The main study of this paper is to focus on the reconstruction of the general shape of the upper Pleistocene hominids, and to examine the cause of cranial changes from the late upper Paleolithic to Mesolithic period in Korea. Biomechanical principles are also applied to reconstruct the social activity of the upper Paleolithic man in Korea. In addition, it is assumed that main proponent of Paleolithic cultures in Korea might be Homo sapiens rather than any other species. Based on the anatomical characteristics of skull, the axillary border of the scapular and the midshaft of the femur, general appearance of Homo sapiens during the Pleistocene in Korea could be reconstructed.     

Patterns of humeral asymmetry among Late Pleistocene humans

Human humeral diaphyseal asymmetry in midshaft and mid-distal rigidity is assessed through the Late Pleistocene in samples of late archaic (Neandertal) and early modern humans. It is considered with respect to directionality (handedness), levels of asymmetry, body size and sexual differences. The overall Late Pleistocene sample indicates a right-handed preference in frequencies (right: 74.8%, left: 15.0%, ambiguous: 10.3%), which are similar to those of recent human samples. Average levels of humeral asymmetry are elevated relative to Holocene samples through all but the small Middle Paleolithic modern human and eastern Eurasian late Upper Paleolithic samples. Humeral asymmetry is especially high among the males relative to the females, and the possibility of a division of labor between uni-manual tasks (mostly male) and bi-manual tasks (mostly female) is considered. At the same time, there is a general pattern of increased asymmetry with larger body size, but it remains unclear to what extent it reflects body size versus sexual effects on bilateral humeral loading. There do not appear to have been substantial changes in humeral asymmetry through time, indicating a continuity of similar manual behavioral patterns through the Late Pleistocene, despite considerable changes in technology through the Late Pleistocene.     

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]     

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.     

Les relations entre l’Extrême-Orient eurasien et le nord de l’archipel du Japon au Paléolithique

The aim of this paper is to present a review of current knowledge concerning the Paleolithic records and the related natural environmental setting in the Eurasian Far East and Hokkaido, located at the northern tip of the Japanese islands. At present, it is quite difficult to answer whether the archaic humans dispersed from Siberia and northern China across the Amur River basin and Sakhalin into Hokkaido or not, because there is no reliable evidence indicating the Lower and Middle Paleolithic in Hokkaido. We demonstrate that the Upper Paleolithic assemblages in Hokkaido can be divided into at least three phases such as the early Upper Paleolithic (EUP), the middle Upper Paleolithic (MUP), and the late Upper Paleolithic (LUP), based on a synthesis of available radiocarbon dates and the techno-typological characteristics of lithic assemblages. It is reasonable to suggest that the lithic assemblage from the Rubenosawa site, located in northern Hokkaido, and some of lithic assemblages at the transition from the Middle to Upper Paleolithic or the initial Upper Paleolithic (IUP) in Siberia share the relative similarities of techno-typological attributes in the reduction sequences, although the reliable radiocarbon dates have not been obtained from the Rubenosawa site unfortunately. Also, the emergence of microblade technology at the MUP in Hokkaido, such as represented by the microblade assemblage recovered from the Kashiwadai-1 site, central Hokkaido, indicates a close interaction between the Eurasian Far East and Hokkaido. As a result, the comparison of archaeological evidence in these regions provides us with a suggestion that the appearance and development of the Upper Paleolithic assemblages in Hokkaido were sometimes associated with the human dispersions and the mutual contacts crossing between the Eurasian Far East and Hokkaido.     

辽宁建平古人类肱骨形态结构分析

1957年,在辽宁省建平县发现了一根古人类肱骨化石,编号PA103。通过同一批龙骨中筛选的哺乳动物化石,吴汝康推断PA103应该为更新世晚期古人类,并对该化石进行了表面形态特征观察和描述。为了对PA103化石的内外结构进行更全面的了解,除了线性测量数据的对比,本文还通过计算机断层扫描技术,结合生物力学和形态示量图分析对建平古人类右侧肱骨化石PA103进行了分析。通过本研究发现,PA103骨干横断面的生物力学粗壮度和力学形状指数明显小于尼安德特人,而与同时期欧亚大陆古人类不利手侧最为接近,这说明建平人右侧肱骨可能不是惯用手,同时,建平人的行为活动应该与同时期同地区的古人类处于同一水平,而小于尼安德特人。整体来看,PA103骨干骨密质厚度和截面惯性矩与近现代人的分布模式较为接近,除局部数值增大外,其整体数值小于近现代人的平均水平,这可能与遗传或行为活动有关,由于缺少古人类化石对比数据,更详细的了解还需后期开展更多相关的研究。     

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.     

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.     

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.     

Late pleistocene human femoral diaphyseal curvature

Anterior femoral curvature is a consistent characteristic of Pleistocene and recent humans, although variation exists in the degree of curvature among individuals and across populations. In particular, one group, the Neandertals, has been characterized for a century as having marked femoral curvature. To evaluate the degree of anterior femoral curvature in both Neandertals and other Late Pleistocene humans, their curvature subtenses and proximodistal positions were evaluated in the context of recent human variation. Recent human comparisons show little relationship between subtense (absolute curvature) and femoral length, suggesting that an index that incorporates subtense relative to the length of the femur is inappropriate for between-group assessments. Neandertals were statistically indistinguishable from Middle or earlier Upper Paleolithic modern humans in the degree of absolute curvature, all of whom had greater curvature on average than all later humans. Additionally, Neandertals and Qafzeh-Skhul early modern humans had a more distal point of maximum curvature than any other group. Curvature was not strongly correlated with functional considerations including body mass estimates, surrogate variables for body size, proximal femoral articular orientation, or knee anteroposterior dimensions. The functional role of femoral anterior curvature is unknown; however, the general decrease in curvature subtense closely parallels the between-group changes in inferred levels of mobility from femoral diaphyseal robusticity and shape, suggesting that femoral curvature may reflect mobility levels and patterns among Late Pleistocene and recent humans.     

Biomechanical approach to the reconstruction of activity patterns in Neolithic Western Liguria, Italy

This paper investigates the changes in upper and lower limb robusticity and activity patterns that accompanied the transition to a Neolithic subsistence in western Liguria (Italy). Diaphyseal robusticity measures were obtained from cross-sectional geometric properties of the humerus and femur in a sample of 16 individuals (eight males and eight females) dated to about 6,000-5,500 BP. Comparisons with European Late Upper Paleolithics (LUP) indicate increased humeral robusticity in Neolithic Ligurian (NEOL) males, but not in females, with a significant reduction in right-left differences in both sexes. Sexual dimorphism in robusticity increases in upper and lower limb bones. Regarding the femur, while all female indicators of bending strength decrease steadily through time, values for NEOL males approach those of LUP. This suggests high, and unexpected, levels of mechanical stress for NEOL males, probably reflecting the effects of the mountainous terrain on lower limb remodeling. Comparisons between NEOL males and a small sample of LUP hunter-gatherers from the same area support this interpretation. In conclusion, cross-sectional geometry data indicate that the transition to Neolithic economies in western Liguria did not reduce functional requirements in males, and suggest a marked sexual division of labor involving a more symmetrical use of the upper limb, and different male-female levels of locomotory stress. When articulated with archaeological, faunal, paleopathological, and ethnographic evidence, these results support the hypothesis of repetitive, bimanual use of axes tied to pastoral activities in males, and of more sedentary tasks linked to agriculture in females.