The oldest anatomically modern humans from far southeast Europe: direct dating, culture and behavior

Background

Anatomically Modern Humans (AMHs) are known to have spread across Europe during the period coinciding with the Middle to Upper Paleolithic transition. Whereas their dispersal into Western Europe is relatively well established, evidence of an early settlement of Eastern Europe by modern humans are comparatively scarce.

Methodology/Principal Finding

Based on a multidisciplinary approach for the study of human and faunal remains, we describe here the oldest AMH remains from the extreme southeast Europe, in conjunction with their associated cultural and paleoecological background. We applied taxonomy, paleoecology, and taphonomy combined with geomorphology, stratigraphy, archeology and radiocarbon dating. More than 160 human bone remains have been discovered. They originate from a well documented Upper Paleolithic archeological layer (Gravettian cultural tradition) from the site of Buran-Kaya III located in Crimea (Ukraine). The combination of non-metric dental traits and the morphology of the occipital bones allow us to attribute the human remains to Anatomically Modern Humans. A set of human and faunal remains from this layer has been radiocarbon dated by Accelerator Mass Spectrometry. The direct-dating results of human bone establish a secure presence of AMHs at 31,900+240/−220 BP in this region. They are the oldest direct evidence of the presence of AMHs in a well documented archeological context. Based on taphonomical observations (cut marks and distribution of skeletal elements), they represent the oldest Upper Paleolithic modern humans from Eastern Europe, showing post-mortem treatment of the dead as well.

Conclusion/Significance

These findings are essential for the debate on the spread of modern humans in Europe during the Upper Paleolithic, as well as their cultural behaviors.     

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.     

The early upper Paleolithic of eastern Europe reconsidered

Artifacts of Paleolithic age were first recognized in eastern Europe during the 1870s. Archeologists have struggled ever since to integrate them into the better known record of western Europe, where the interpretive framework of Paleolithic archeology was originally developed. The essential elements of both the Middle and Upper Paleolithic were recognized quickly in eastern Europe, and a close connection with a major middle Upper Paleolithic industry of central Europe (Gravettian) was established many years ago. The early Upper Paleolithic (EUP) has remained a major challenge, however; it is represented primarily by a bewildering array of local archeological cultures that exhibit limited similarity to contemporaneous industries of western and central Europe.6-9.     

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.     

The beginning of the Upper Paleolithic in the Iranian Zagros. A taphonomic approach and techno-economic comparison of Early Baradostian assemblages from Warwasi and Yafteh (Iran)

Southwest Asia is a key region in current debates surrounding the appearance of the first cultures attributed to anatomically modern humans, particularly the Aurignacian and preceding cultural units of the Iranian Zagros, Levant, and the Balkans (Baradostian, Ahmarien, Kozarnikien, etc.). The Zagros mountain range encompasses an immense territory that remains understudied with regard to the Upper Paleolithic as well as the first bladelet industries traditionally presumed to be the work of anatomically modern humans. Concerning the emergence of the Aurignacian, the sites of Warwasi rockshelter and Yafteh cave in the central Zagros are considered to show evidence of in situ evolution of the Upper Paleolithic from the local Mousterian. This hypothesis is tested by way of a taphonomic, techno-typological and economic approach applied to the Upper Paleolithic levels of Warwasi (spits LL–AA) and Yafteh (the series from the lower part of the sequence). A comparison of the techno-economic features of both assemblages demonstrates a conceptual bond with contemporaneous techno-complexes from Levant and Europe (Ahmarian, Protoaurignacian, etc.). The techno-typological Middle Paleolithic character of the Warwasi lithic assemblage permits a discussion of a possible in situ dependence/continuum from the Mousterian or perhaps particular activities linked to the type of the occupation of the site. However, bladelet technology cannot be considered as rooted in the Zagros Mousterian. Consequently the origin of the Aurignacian sensu stricto has to be reconsidered.     

L’impact de l’environnement sur l’évolution biologique et culturelle dans les périodes de transition en Préhistoire : Paléolithique ancien/moyen et Paléolithique moyen/supérieur

This paper deals with the environmental changes during two important transitional periods, namely Lower/Middle Paleolithic and Middle/Upper Paleolithic transitions, and with their impact on biological and cultural evolutions, in the context of central, eastern and northwestern Europe, where ice-sheet transgressions induced particular paleogeographical and paleoecological conditions. Another country was studied too, the Balkans, a privileged area for bilateral contacts between Europe and the Near East.     

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.     

Timescales and cultural process at 40,000 BP in the light of the Campanian Ignimbrite eruption, Western Eurasia

Significant new information shows that the Campanian Ignimbrite (CI) eruption from the Phlegrean Fields, southern Italy, was much larger than hitherto supposed and in fact one of the largest late Quaternary explosive events. The eruption can be dated to 40,000 calendar years ago, within the interval of the so-called Middle to Upper Paleolithic 'transition'. Its position can be precisely correlated with a number of other environmental events, including Heinrich Event 4 (HE4), the Laschamp excursion, and a particular cosmogenic nuclide peak. In view of this unique combination of factors, we studied the CI volcanic catastrophe with particular attention to its impact on climate and human ecosystems, including potential interference with ongoing processes of cultural evolution (biological evolution is best left aside for the moment). The contribution of this research is chronological and ecological. The CI volcanic event provides an unequalled means of correlating stratigraphic sequences across Western Eurasia, either directly or indirectly, and affords a unique opportunity to establish the age and climatic context of important archaeological sequences. Ecologically, the CI eruption inevitably interacted with the beginning of HE4 in terms of atmospheric feedback systems. Their combined forcing produced a sudden and at least hemispheric climatic deterioration; a 'volcanic winter' scenario cannot be ruled out. Paleolithic occupation was severely altered throughout the direct-impact zone of the eruption and likely along fringe areas in southern and southeastern Europe. The above observations call for a reconsideration of the processes and rhythms involved in the Middle to Upper Paleolithic 'transition'. A tentative model is suggested that links the exceptional environmental stress at 40,000 BP with processes already active in Paleolithic societies, leading to a period of accelerated change in cultural configurations. These eventually evolved into an Upper Paleolithic proper at a later date. The evidence to invoke allochthonous cultural input or invasionist scenarios is not considered compelling.     

Nonhuman primate hybridization and the taxonomic status of Neanderthals

The present study examines the taxonomic status of Middle Paleolithic Neanderthals by comparing their observed minimum genetic divergence from Upper Paleolithic modern humans in Europe with that observed between macaque species from Sulawesi that are known to hybridize and fully intergrade in the wild. The genetic divergence, and differentiation between Neanderthals and Upper Paleolithic modern humans, as indicated by pairwise minimum genetic distances and F(ST) values calculated from the estimated minimum genetic relationship (R) matrix derived from craniometric data, are significantly greater than those observed both between hybridizing and noninterbreeding Sulawesi macaque species, suggesting that mate recognition and the possibility of gene flow between Neanderthals and Upper Paleolithic modern humans might have been greatly reduced. These results support a species-level taxonomic distinction for the Neanderthals as suggested by proponents of the replacement model. Furthermore, assumptions regarding the monophyletic origin of modern humans from outside Europe are likely valid.     

Early Upper Paleolithic Man and Late Middle Paleolithic Tools

The appearance of anatomically modern Homo sapiens in Europe, the Near East, and Africa must represent either an in situ evolution of Neandertals or a migration. Those who suggest the latter claim a sudden replacement of Neandertals by anatomically modern Homo sapiens. However, the "evidence" actually cited claims only the sudden replacement of Middle by Upper Paleolithic industries. We criticize the migration explanation on two grounds. (1) There is no "sudden replacement" of Middle Paleolithic by Upper Paleolithic industries, but rather a gradual change in the frequencies of already present tools. Numerous sites in these areas exhibit transitional industries. (2) Concomitantly, there is no morphological evidence indicating a "sudden replacement" of hominids. There is no absolute association between anatomically modern Homo sapiens and Upper Paleolithic industries. Instead, the evidence clearly shows that early anatomically modern Homo sapiens is a late Middle Paleolithic local phenomenon .     

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.     

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.     

Radiocarbon dating the late Middle Paleolithic and the Aurignacian of the Swabian Jura

Many lines of evidence point to the period between roughly 40 and 30 ka BP as the period in which modern humans arrived in Europe and displaced the indigenous Neandertal populations. At the same time, many innovations associated with the Upper Paleolithic--including new stone and organic technologies, use of personal ornaments, figurative art, and musical instruments--are first documented in the European archaeological record. Dating the events of this period is challenging for several reasons. In the period about six to seven radiocarbon half-lives ago, variable preservation, pre-treatment, and sample preparation can easily lead to a lack of reproducibility between samples and laboratories. A range of biological, cultural, and geological processes can lead to mixing of archaeological strata and their contents. Additionally, some data sets point to this period as a time of significant spikes in levels of atmospheric radiocarbon. This paper assesses these questions in the context of the well-excavated and intensively studied caves of Geissenkl?sterle and Hohle Fels in the Swabian Jura of southwestern Germany. We conclude that variable atmospheric radiocarbon production contributes to the problems of dating the late Middle Paleolithic and the early Upper Paleolithic. To help establish a reliable chronology for the Swabian Aurignacian, we are beginning to focus our dating program on short-lived, stratigraphically secure features to see if they yield reproducible results. This approach may help to test competing explanations for the noisy and often non-reproducible results that arise when trying to date the transition from the Middle to the Upper Paleolithic.     

A modern human humerus from the early aurignacian of Vogelherdhöhle (Stetten, Germany)

Implicit in much of the discussion of the cultural and population biological dynamics of modern human origins in Europe is the assumption that the Aurignacian, from its very start, was made by fully modern humans. The veracity of this assumption has been challenged in recent years by the association of Neandertal skeletal remains with a possibly Aurignacian assemblage at Vindija Cave (Croatia) and the association of Neandertals with distinctly Upper Paleolithic (but non-Aurignacian) assemblages at Arcy-sur-Cure and St. C?esaire (France). Ideally we need human fossil material that can be confidently assigned to the early Aurignacian to resolve this issue, yet in reality there is a paucity of well-provenanced human fossils from early Upper Paleolithic contexts. One specimen, a right humerus from the site of Vogelherd (Germany), has been argued, based on its size, robusticity, and muscularity, to possibly represent a Neandertal in an Aurignacian context. The morphological affinities of the Vogelherd humerus were explored by univariate and multivariate comparisons of humeral epiphyseal and diaphyseal shape and strength measures relative to humeri of Neandertals and Early Upper Paleolithic (later Aurignacian and Gravettian) modern humans. On the basis of diaphyseal cross-sectional geometry, deltoid tuberosity morphology, and distal epiphyseal morphology, the specimen falls clearly and consistently with European early modern humans and not with Neandertals. Along with the other Vogelherd human remains, the Vogelherd humerus represents an unequivocal association between the Aurignacian and modern human morphology in Europe.     

李家沟遗址的石器工业

本文介绍河南新密李家沟遗址发现的石制品, 并进行简要讨论。该遗址包含了从旧石器时代晚期一直到新石器时代多个文化阶段的遗存。石器工业在不同文化阶段表现出不同的特点。除了旧石器晚期较早阶段的石片石器组合和旧石器时代晚期之末的细石器组合之外, 还在新石器早期文化乃至裴李岗文化阶段的文化遗存中发现数量较多的打制石器。其中最值得关注的是与典型细石器共存的磨刃石锛与陶片。通过对石制品的初步观察可知, 打制石器并不只存在于旧石器时代, 而是延续到新石器早期甚至可能裴李岗文化时期, 打制石器仍然在继续使用。这种情况说明, 李家沟以及中原地区旧、新石器时代的石器工具的变化过渡是一种逐渐变化的过程。     

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。     

Who made the Aurignacian and other early Upper Paleolithic industries?

The Aurignacian is typically taken as a marker of the spread of anatomically modern humans into Europe. However, human remains associated with this industry are frustratingly sparse and often limited to teeth. Some have suggested that Neandertals may, in fact, be responsible for the Aurignacian and the earliest Upper Paleolithic industries. Although dental remains are frequently considered to be taxonomically undiagnostic in this context, recent research shows that Neandertals possess a distinct dental pattern relative to anatomically modern humans. Even so, it is rare to find mandibles or maxillae that preserve all or most of their teeth; and, the probability of correctly identifying individuals represented by only a few teeth or a single tooth is unknown. We present a Bayesian statistical approach to classifying individuals represented exclusively by teeth into two possible groups. The classification is based on dental trait frequencies and sample sizes for ‘known’ samples of 95 Neandertals and 63 Upper Paleolithic modern humans. In a cross validation test of the known samples, 89% of the Neandertals and 89% of the Upper Paleolithic modern humans were classified correctly. We then classified an ‘unknown’ sample of 52 individuals: 34 associated with Aurignacian or other (non-Châtelperronian) early Upper Paleolithic industries, 15 associated with the Châtelperronian, and three unassociated. Of the 34 early Upper Paleolithic-associated individuals, 29 were assigned to modern humans, which is well within the range expected (95% of the time 26-33) with an 11% misclassification rate for an entirely modern human sample. These results provide some of the strongest evidence that anatomically modern humans made the Aurignacian and other (non-Châtelperronian) early Upper Paleolithic industries.     

Seasonal variations of the middle-upper paleolithic transition at El castillo, Cueva Morín and El pendo (Cantabria, Spain)

With debate escalating in regard to the prolonged contemporaneity of neandertal and modern human groups in the Franco-Cantabrian region on the one hand, and the late persistence of neandertals (until ca. 28-30,000 B.P.) and Mousterian industries in southern Iberia on the other; sites with Mousterian-Upper Paleolithic sequences from northern Spain play a pivotal role in the ongoing investigation of the Middle-Upper Paleolithic transition in western Europe. An important line of inquiry into the nature of social and economic change from the Middle to Upper Paleolithic is the monitoring of shifts in land use and resource procurement patterns. The recognition of short-term, seasonal patterning in settlement and resource provisioning may provide insights into changes in mobility, territoriality, and social organization that might otherwise be missed. This paper presents results of a seasonality study of fauna from archaeological levels spanning the Middle-Upper Paleolithic transition from the sites of El Castillo, El Pendo, and Cueva Morín in Cantabrian Spain. Data concerning season of death and age at death of prey animals presented here are derived from dental growth mark (increment, annuli) analysis. These data, along with other artifactual and faunal evidence suggest to us that: (1) economic strategies and technologies pervasive in the Upper Paleolithic are rooted in the Cantabrian Middle Paleolithic; and, (2) the apparent increase in deposits from the Middle through Upper Paleolithic may be the signature of a gradual increase in logistical economic strategies including the heightened level of social organization required for their implementation.     

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.