Reassessment of TL age estimates of burnt flints from the Paleolithic site of Tabun Cave, Israel

The stratigraphy of Tabun Cave (Mt. Carmel), which comprises one of the longest sequences of Lower and Middle Paleolithic of the Near East, is widely used as a reference in debates on the evolution of Paleolithic industries and on the origin of modern humans and their relationship to the Neandertals. Considering the methodological improvements during the last ten years, the thermoluminescence (TL) dates of heated flints frequently quoted in the literature require an update. New TL results are discussed and compared with radiometric data recently obtained for this site, in particular by the ESR method, and with those obtained for other Levantine sites. The chronological framework previously proposed for the Paleolithic industries of the area is then re-examined.     

Time for the Middle to Upper Paleolithic transition in Europe

The Middle to Upper Paleolithic transition is a key period of change in the prehistory of the Old World and one of the most studied issues in paleoanthropology, as the nature of the transition(s) is still, after at least a century of archaeological research, largely unknown. Many of the issues at stake in the transition relate to the problem of building a reliable chronology for this period, which is at the limits of the radiocarbon method. The papers in this volume show that much progress has been made in our chronological knowledge of significant aspects of the transition, such as the age of the most recent Neandertal fossils and the earliest modern human remains in Europe, and the inferred overlap between the Chatelperronian and the Aurignacian. At the same time, the volume also shows where the chronological database for the period 40 to 30 ka 14C BP is flawed and that significant contextual and methodological problems have been underestimated in a number of studies of the biological and cultural changes during this crucial period. Chronology is employed by paleoanthropologists to relate the record of the Middle to Upper Paleolithic transition to major biological and cultural developments. This paper takes a broader paleoanthropological perspective and attempts to evaluate and, to some degree, synthesize the main results of these proceedings, while also presenting a brief discussion of the Middle and Upper Paleolithic archaeological and fossil records, and possible explanations for the differences between the two, focusing on the role of differences in the ecology of Neandertals and early European modern humans.     

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.     

New data on the late Neandertals: Direct dating of the Belgian Spy fossils

In Eurasia, the period between 40,000 and 30,000 BP saw the replacement of Neandertals by anatomically modern humans (AMH) during and after the Middle to Upper Paleolithic transition. The human fossil record for this period is very poorly defined with no overlap between Neandertals and AMH on the basis of direct dates. Four new ¹⁴C dates were obtained on the two adult Neandertals from Spy (Belgium). The results show that Neandertals survived to at least ≈36,000 BP in Belgium and that the Spy fossils may be associated to the Lincombian–Ranisian–Jerzmanowician, a transitional techno‐complex defined in northwest Europe and recognized in the Spy collections. The new data suggest that hypotheses other than Neandertal acculturation by AMH may be considered in this part of Europe. Am J Phys Anthropol, 2009. © 2008 Wiley‐Liss, Inc.     

Throwing in the Middle and Upper Paleolithic: inferences from an analysis of humeral retroversion

When in evolutionary history did long-range projectile weapons become an important component of hunting toolkits? The archeological evidence for the development of projectile weaponry is complex and generally indirect, and has led to different conclusions about the origin and spread of this technology. Lithic evidence from the Middle Stone Age (MSA) has led some researchers to suggest that true long- range projectile weaponry developed in Africa perhaps as early as 80,000 years ago, and was part of the subsistence toolkit carried by modern humans who expanded out of Africa after 50,000 years ago. Alternatively, temporal patterns in the morphology of pointed lithics has led others to posit an independent, convergent origin of projectile weaponry in Africa, the Near East, and Europe during the interval between 50,000-40,000 years ago. By either scenario, projectile weapons would not have been a component of the hunting arsenal of Neandertals, but may have been in use by European early modern humans and thus, projectile technology may have entered into the competitive dynamics that existed between these two groups. The origins of projectile weapons can be addressed, in part, through analyses of the skeletal remains of the prehistoric humans who made and used them. Habitual behavior patterns—including those related to the production and use of technology—can be imprinted on the skeleton through both genetic and epigenetic pathways. Recent studies in the field of sports medicine indicate that individuals who engage in habitual throwing have increased humeral retroversion angles in their throwing arms and a greater degree of bilateral asymmetry in retroversion angles than do non-throwers. This contribution investigates humeral torsion through analysis of the retroversion angle in samples of Eurasian Neandertals, European early modern humans of the middle and late Upper Paleolithic, and comparative samples of recent humans. This analysis was conducted under the assumption that if throwing-based projectile weaponry was used by early modern Europeans but not Neandertals, Upper Paleolithic samples should be similar to recent human groups engaged in habitual throwing in the degree of humeral retroversion in the dominant limb and in bilateral asymmetry in this feature. Neandertals on the other hand, would not be expected to show marked asymmetry in humeral retroversion. Consistent with other studies, Neandertals exhibit increased retroversion angles (decreased humeral torsion or a more posteriorly oriented humeral head) relative to most modern human samples, although this appears more likely related to body form and overall activity levels than to habitual throwing. Although Neandertals with bilaterally preserved humeri sufficient for measurement are rare (consisting of only two males and one female), levels of bilateral asymmetry in humeral retroversion are low, suggesting a lack of regular throwing. While patterning across fossil and comparative samples in levels of humeral retroversion was not clear cut, males of both the middle and late Upper Paleolithic demonstrate a high level of bilateral asymmetry, comparable to or in excess of that seen in samples of throwing athletes. This may indicate habitual use of throwing-based projectile weaponry by middle Upper Paleolithic times. Small sample sizes and relatively great variance in the fossil samples makes these results, however, suggestive rather than conclusive.     

现代人出现和演化的化石证据

近年来,新材料的发现、新测年结果的更新以及分子生物学的加入,刷新了关于现代人在各主要区域出现时间、在迁徙扩散中与古老人群的交流模式、晚更新世晚期现代人演化复杂场景等方面的认识。本文梳理了关于现代人出现和演化路径的主要化石证据和研究成果。目前化石证据显示：非洲准现代人的出现时间最早可到MIS 9阶段,非洲现代人在中更新世晚期到晚更新世化石证据连续;欧亚大陆现代人的出现时间可追溯到MIS 6阶段,在MIS 5a-MIS 4阶段经历“瓶颈期”,在MIS 3阶段开始广泛分布。从已有证据来看,MIS 3和MIS 2阶段,欧亚大陆不同区域的现代人演化链条并非单一线性,而是呈现有断点的“网状”演化模式。     

Neandertals and moderns mixed,and it matters

Twenty‐five years ago, the Middle‐to‐Upper Paleolithic transition in Europe could be represented as a straightforward process subsuming both the emergence of symbolic behavior and the replacement of Neandertals by modern humans. The Aurignacian was a proxy for the latter, during which enhanced cognitive capabilities explained ornaments and art. The few instances of Neandertal symbolism were deemed to long postdate contact and dismissed as “imitation without understanding,” if not geological contamination. Such views were strengthened by the recent finding that, in southern Africa, several features of the European Upper Paleolithic, including bone tools, ornaments, and microliths, emerged much earlier. Coupled with genetic suggestions of a recent African origin for extant humans, fossil discoveries bridging the transition between “archaics” and “moderns” in the realm of anatomy (Omo‐Kibish, Herto) seemingly closed the case. Over the last decade, however, taphonomic critiques of the archeology of the transition have made it clear that, in Europe, fully symbolic sapiens behavior predates both the Aurignacian and moderns. And, in line with evidence from the nuclear genome rejecting strict replacement models based on mtDNA alone, the small number of early modern specimens that passed the test of direct dating present archaic features unknown in the African lineage, suggesting admixture at the time of contact. In the realm of culture, the archeological evidence also supports a Neandertal contribution to Europe's earliest modern human societies, which feature personal ornaments completely unknown before immigration and are characteristic of such Neandertal‐associated archeological entities as the Châtelperronian and the Uluzzian. The chronometric data suggest that, north of the Ebro divide, the entire interaction process may have been resolved within the millennium centered around 42,000 calendar years ago. Such a rapid absorption of the Neandertals is consistent with the size imbalance between the two gene reservoirs and further supports significant levels of admixture.     

Genetic evidence and the modern human origins debate

A continued debate in anthropology concerns the evolutionary origin of 'anatomically modern humans' (Homo sapiens sapiens). Different models have been proposed to examine the related questions of (1) where and when anatomically modern humans first appeared and (2) the genetic and evolutionary relationship between modern humans and earlier human populations. Genetic data have been increasingly used to address these questions. Genetic data on living human populations have been used to reconstruct the evolutionary history of the human species by considering how global patterns of human variation could be produced given different evolutionary scenarios. Of particular interest are gene trees that reconstruct the time and place of the most recent common ancestor of humanity for a given haplotype and the analysis of regional differences in genetic diversity. Ancient DNA has also allowed a direct assessment of genetic variation in European Neandertals. Together with the fossil record, genetic data provide insight into the origin of modern humans. The evidence points to an African origin of modern humans dating back to 200,000 years followed by later expansions of moderns out of Africa across the Old World. What is less clear is what happened when these early modern humans met preexisting 'archaic human' populations outside of Africa. At present, it is difficult to distinguish between a model of total genetic replacement and a model that includes some degree of genetic mixture.     

Modern hunting behavior in the early Middle Paleolithic: faunal remains from Misliya Cave, Mount Carmel, Israel

Understanding the behavioral adaptations and subsistence strategies of Middle Paleolithic humans is critical in the debate over the evolution and manifestations of modern human behavior. The study of faunal remains plays a central role in this context. Until now, the majority of Levantine archaeofaunal evidence was derived from late Middle Paleolithic sites. The discovery of faunal remains from Misliya Cave, Mount Carmel, Israel (>200 ka), allowed for detailed taphonomic and zooarchaeological analyses of these early Middle Paleolithic remains. The Misliya Cave faunal assemblage is overwhelmingly dominated by ungulate taxa. The most common prey species is the Mesopotamian fallow deer (Dama mesopotamica), followed closely by the mountain gazelle (Gazella gazella). Some aurochs (Bos primigenius) remains are also present. Small-game species are rare. The fallow deer mortality pattern is dominated by prime-aged individuals. A multivariate taphonomic analysis demonstrates (1) that the assemblage was created solely by humans occupying the cave and was primarily modified by their food-processing activities; and (2) that gazelle carcasses were transported complete to the site, while fallow deer carcasses underwent some field butchery. The new zooarchaeological data from Misliya Cave, particularly the abundance of meat-bearing limb bones displaying filleting cut marks and the acquisition of prime-age prey, demonstrate that early Middle Paleolithic people possessed developed hunting capabilities. Thus, modern large-game hunting, carcass transport, and meat-processing behaviors were already established in the Levant in the early Middle Paleolithic, more than 200 ka ago.     

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.     

Modern Human Origins in the Levant and Western Asia: The Fossil and Archeological Evidence

Upper Pleistocene human fossil and archeological evidence from the Levant and western Asia indicates continuity over the Middle/Upper Paleolithic transition and the transition from archaic to modern Homo sapiens. It also indicates that these transitions did not coincide with each other in time. Both data sets are examined in light of recent claims from molecular biology invoked by replacement advocates in support of the position that morphologically modern humans develop (or arrive) first in the Levant, coexist with archaic Homo sapiens, and then displace or extirpate archaic Homo sapiens. Replacement models that assume no admixture are not supported by the archeological or the human paleontological evidence. Those who would argue for replacement without gene flow must show how it could have occurred without leaving traces of disjunction in the typological and technological aspects of Levantine archeological assemblages, in those aspects of the archeofaunal record that monitor subsistence, and in the evidence from settlement pattern studies.     

The Date of Interbreeding between Neandertals and Modern Humans

Comparisons of DNA sequences between Neandertals and present-day humans have shown that Neandertals share more genetic variants with non-Africans than with Africans. This could be due to interbreeding between Neandertals and modern humans when the two groups met subsequent to the emergence of modern humans outside Africa. However, it could also be due to population structure that antedates the origin of Neandertal ancestors in Africa. We measure the extent of linkage disequilibrium (LD) in the genomes of present-day Europeans and find that the last gene flow from Neandertals (or their relatives) into Europeans likely occurred 37,000–86,000 years before the present (BP), and most likely 47,000–65,000 years ago. This supports the recent interbreeding hypothesis and suggests that interbreeding may have occurred when modern humans carrying Upper Paleolithic technologies encountered Neandertals as they expanded out of Africa.     

The Middle/Later Stone Age transition and cultural dynamics of late Pleistocene East Africa

The Middle to Later Stone Age (MSA/LSA) transition is a prominent feature of the African archeological record that began in some places ~30,000–60,000 years ago, historically associated with the origin and/or dispersal of “modern” humans. Unlike the analogous Middle to Upper Paleolithic transition in Eurasia and associated Neanderthal extinction, the African MSA/LSA record remains poorly documented, with its potential role in explaining changes in the behavioral diversity and geographic range of Homo sapiens largely unexplored. I review archeological and biogeographic data from East Africa, show regionally diverse pathways to the MSA/LSA transition, and emphasize the need for analytical approaches that document potential ancestor‐descendent relationships visible in the archeological record, needed to assess independent invention, population interaction, dispersal, and other potential mechanisms for behavioral change. Diversity within East Africa underscores the need for regional, rather than continental‐scale narratives of the later evolutionary history of H. sapiens.     

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 .     

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.     

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.     

Dental remains from the Grotte du Renne at Arcy-sur-Cure (Yonne)

Human remains associated with the earliest Upper Paleolithic industries are sparse. What is preserved is often fragmentary, making it difficult to accurately assign them to a particular species. For some time it has been generally accepted that Neandertals were responsible for the Chatelperronian and anatomically modern humans for the early Aurignacian industries. However, the recent re-dating of several of the more-complete modern human fossils associated with the early Aurignacian (e.g., Vogelherd) has led some to question the identity of the makers and the context of these early Upper Paleolithic industries. The Grotte du Renne at Arcy-sur-Cure, France has yielded many hominin remains, from Mousterian, Chatelperronian, Aurignacian, and Gravettian layers. Previously, a child's temporal bone from the Chatelperronian Layer Xb was recognized as belonging to a Neandertal; however, most of the teeth from Chatelperronian layers VIII-X remain unpublished. We describe the dental remains from the Chatelperronian layers, place them in a comparative (Mousterian Neandertal and Upper Paleolithic modern human) context, and evaluate their taxonomic status. The teeth (n = 29) represent a minimum of six individuals aged from birth to adult. The permanent dental sample (n = 15) from the Chatelperronian layers of Arcy-sur-Cure exhibits traits (e.g., lower molar mid-trigonid crest) that occur more frequently in Neandertals than in Upper Paleolithic modern humans. Furthermore, several teeth show trait combinations, including Cusp 6/mid-trigonid crest/anterior fovea in the lower second molar, that are rare or absent in Upper Paleolithic modern humans. The deciduous teeth (n = 14) significantly increase the sample of known deciduous hominin teeth and are more similar to Mousterian Neandertals from Europe and Asia than to Upper Paleolithic modern humans. Thus, the preponderance of dental evidence from the Grotte du Renne strongly supports that Neandertals were responsible for the Chatelperronian industry at Arcy-sur-Cure.     

Late archaic and modern Homo sapiens from Europe, Africa, and Southwest Asia: Craniometric comparisons and phylogenetic implications

The craniometric affinities among Neandertals. Upper Palcolithic Europeans, early anatomically modern Southwest Asians, and archaic and modern Africans are investigated using univariate and multivariate methods. For the first time, it is possible to analyse the North African finds Dar-es-Soltane 5, Nazlet Khater, and Wadi Kubbaniya. It was not possible to include the Neandertals from Central Europe due to their poor state of preservation. The results point to, first, a basic distinction between Neandertals on the one hand and modern humans from all geographic regions on the other, and, secondly, to great similarities between modern African and European populations. Late archaic sapiens specimens from Africa were more similar to Upper Paleolithic Europeans than were the Neandertals. The results do not support the hypothesis that a regional evolution giving rise to modern humans took place in Europe. The results are, however, consistent with the hypothesis that modern populations originated in Africa and spread to Europe from there.     

Comparative morphometrics of the primate apical tuft

The relationship between the structure and function of the primate apical tuft is poorly understood. This study addresses several hypotheses about apical tuft morphology using a large modern primate comparative sample. Two indices of tuft size are employed: expansion and robusticity. First, comparisons of relative apical tuft size were drawn among extant nonhuman primate groups in terms of locomotion and phylogenetic category. Both of these factors appear to play a role in apical tuft size among nonhuman primates. Suspensory primates and all platyrrhines had the smallest apical tufts, while terrestrial quadrupeds and all strepsirrhines (regardless of locomotor category) had the largest tufts. Similarly, hypotheses regarding the apical tufts of hominins, especially the large tufts of Neandertals were addressed using a comparison of modern warm- and cold-adapted humans. The results showed that cold-adapted populations possessed smaller apical tufts than did warm-adapted groups. Therefore, the cold-adaptation hypothesis for Neandertal distal phalangeal morphology is not supported. Also, early modern and Early Upper Paleolithic humans had apical tufts that were significantly less expanded and less robust than those of Neandertals. The hypothesis that a large apical tuft serves as support for an expanded digital pulp is supported by radiographic analysis of modern humans in that a significant correlation was discovered between the width of the apical tuft and the width of the pulp. The implications of these findings for hypotheses about the association of apical tuft size and tool making in the hominin fossil record are discussed.