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.     

The reality of Neandertal symbolic behavior at the Grotte du Renne, Arcy-sur-Cure, France

Background

The question of whether symbolically mediated behavior is exclusive to modern humans or shared with anatomically archaic populations such as the Neandertals is hotly debated. At the Grotte du Renne, Arcy-sur-Cure, France, the Châtelperronian levels contain Neandertal remains and large numbers of personal ornaments, decorated bone tools and colorants, but it has been suggested that this association reflects intrusion of the symbolic artifacts from the overlying Protoaurignacian and/or of the Neandertal remains from the underlying Mousterian.

Methodology/Principal Findings

We tested these hypotheses against the horizontal and vertical distributions of the various categories of diagnostic finds and statistically assessed the probability that the Châtelperronian levels are of mixed composition. Our results reject that the associations result from large or small scale, localized or generalized post-depositional displacement, and they imply that incomplete sample decontamination is the parsimonious explanation for the stratigraphic anomalies seen in the radiocarbon dating of the sequence.

Conclusions/Significance

The symbolic artifacts in the Châtelperronian of the Grotte du Renne are indeed Neandertal material culture.     

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.     

Before the transition? The final middle Palaeolithic lithic industry from the Grotte du Renne (layer XI) at Arcy-sur-Cure (Burgundy,France)

The Grotte du Renne at Arcy-sur-Cure (Burgundy, France) has yielded remains from several occupation layers, extending from the end of the middle Palaeolithic to the upper Palaeolithic. The last Mousterian layer (XI), which precedes the Châtelperronian occupations (layers Xa, Xb, Xc, IX, VIII), was previously interpreted as a precursor of the transition complex. The new study of the lithic assemblage of layer XI, in the context of the other final Mousterian occupations in Arcy (Grotte du Bison, Galerie Schoepflin Rotonde, Grotte de l’Hyène) challenges this hypothesis. The Mousterian Arcy complex can be divided into three main techno-complexes: Levallois–Charentian, Discoid–Denticulate and Levallois Mousterian. We observed a rupture between the final Mousterian and Châtelperronian industries as regards all the aspects of the technical system; production methods, blank management, tool types, and some changes in raw material composition.     

A critical review of the German Paleolithic hominin record

We review the hominin fossil record from western Central Europe in light of the recent major revisions of the geochronological context. The mandible from Mauer (Homo heidelbergensis), dated to circa 500,000 years ago, continues to represent the earliest German hominin and may coincide with the occupation of Europe north of the high alpine mountain chains. Only limited new evidence is available for the Middle Pleistocene, mostly in the form of skull fragments, a pattern that may relate to taphonomic processes. These finds and their ages suggest the gradual evolution of a suite of Neandertal features during this period. Despite new finds of classic Neandertals, there is no clear proof for Neandertal burial from Germany. Alternatively, cut marks on a skull fragment from the Neandertal type site suggest special treatment of that individual. New Accelerator Mass Spectrometry (AMS) radiocarbon dates of previous finds leave little reliably dated evidence for anatomically modern humans (AMH) in Europe before 30,000 BP; the remains from Hahn?fersand, Binshof-Speyer, Paderborn-Sande, and Vogelherd are now of Holocene age. Thus, a correlation of AMH with the Aurignacian remains to be proven, and the general idea of a long coexistence of Neandertals and AMH in Europe may be questioned. In western Central Europe, evidence of Gravettian human fossils is also very limited, although a new double grave from lower Austria may be relevant. The only dated burial from the German Upper Paleolithic (from Mittlere Klause) falls into a time period (circa 18,600 BP) represented by only a few occupation sites in western Central Europe. A number of human remains at Magdalenian sites appear to result from variable (secondary) burial practices. In contrast, the Final Paleolithic (circa 12,000-9600 cal. BC) yields an increase of hominin finds, including multiple burials (Bonn-Oberkassel, Neuwied-Irlich), similar to the situation in western and southern 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.     

Neandertal versus Modern Human Dietary Responses to Climatic Fluctuations

The Neandertal lineage developed successfully throughout western Eurasia and effectively survived the harsh and severely changing environments of the alternating glacial/interglacial cycles from the middle of the Pleistocene until Marine Isotope Stage 3. Yet, towards the end of this stage, at the time of deteriorating climatic conditions that eventually led to the Last Glacial Maximum, and soon after modern humans entered western Eurasia, the Neandertals disappeared. Western Eurasia was by then exclusively occupied by modern humans. We use occlusal molar microwear texture analysis to examine aspects of diet in western Eurasian Paleolithic hominins in relation to fluctuations in food supplies that resulted from the oscillating climatic conditions of the Pleistocene. There is demonstrable evidence for differences in behavior that distinguish Upper Paleolithic humans from members of the Neandertal lineage. Specifically, whereas the Neandertals altered their diets in response to changing paleoecological conditions, the diets of Upper Paleolithic humans seem to have been less affected by slight changes in vegetation/climatic conditions but were linked to changes in their technological complexes. The results of this study also indicate differences in resource exploitation strategies between these two hominin groups. We argue that these differences in subsistence strategies, if they had already been established at the time of the first contact between these two hominin taxa, may have given modern humans an advantage over the Neandertals, and may have contributed to the persistence of our species despite habitat-related changes in food availabilities associated with climate fluctuations.     

The Pech-de-l'Azé I Neandertal child: ESR, uranium-series, and AMS 14C dating of its MTA type B context

The Pech-de-l'Azé I skull and mandible are included in the juvenile Neandertal remains from Europe. However, some preserved features in the cranial skeleton seem to distinguish the specimen from other Neandertal children. Unfortunately, the stratigraphic position and dating of this child has never been clear. Our recent work on unpublished archives show that the Pech-de-l'Azé I Neandertal child was discovered at the bottom of layer 6, attributed to the Mousterian of Acheulean tradition type B. These skull and mandible are the first diagnostic human remains (aside from an isolated tooth) attributed to the Mousterian of Acheulian tradition (MTA) type B. Consequently, we confirm that Neandertals were the makers of this Mousterian industry, which is characterized by unusual high frequencies of Upper Paleolithic type tools, elongated blanks and blades. We were able to date the context of the hominid remains by dating layer 6 and the layers above and beneath it using ESR, coupled ESR/(230)Th/(234)U (coupled ESR/U-series), and AMS (14)C. Coupled ESR/U-series results on 16 mammalian teeth constrain the age of the uppermost layer 7 to 41-58ka, and layer 6 to 37-51ka. The wide spread in each age estimate results mainly from uncertainties in the gamma-dose rate. These ages are concordant with AMS (14)C ages of two bones coming from the top of layer 6, which provide dates of about 41.7-43.6ka cal BP. A combination of stratigraphic arguments and dating results for layers 6 and 7 show that the Neandertal child cannot be older than 51ka or younger than 41ka. The lowermost layer 4 is shown to be older than 43ka by the principle of superposition and ESR dating in the immediately overlying layer 5. This study shows that the MTA type B had been manufactured by Neandertals before the arrival of anatomically modern humans in the local region. Additionally, by providing a firm chronological framework for the specific morphometric the features of Pech-de-l'Azé I Neandertal child, this study is a new step toward the understanding of temporal and spatial changes in the ontogenesis of Neandertals in south-western Europe during oxygen isotope stages 5-3.     

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.     

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.     

Did the lateral enamel of Neandertal anterior teeth grow differently from that of modern humans?

The formation of lateral enamel in Neandertal anterior teeth has been the subject of recent studies. When compared to the anterior teeth of modern humans from diverse regions (Point Hope, Alaska; Newcastle upon Tyne, England; southern Africa), Neandertal anterior teeth appear to fall within the modern human range of variation for lateral enamel formation time. However, the lateral enamel growth curves of Neandertals are more linear than those of these modern human samples. Other researchers have found that the lateral enamel growth curves of Neandertals are more linear than those of Upper Paleolithic and Mesolithic modern humans as well. The statistical significance of this apparent difference between Neandertal and modern human lateral enamel growth curves is analyzed here. The more linear Neandertal enamel growth curves result from the smaller percentage of total perikymata located in the cervical halves of their teeth. The percentage of total perikymata in the cervical halves of teeth is therefore compared between the Neandertal sample (n=56 teeth) and each modern human population sample: Inuit (n=65 teeth), southern African (n=114 teeth), and northern European (n=115 teeth). There are 18 such comparisons (6 tooth types, Neandertals vs. each of the three modern human populations). Eighteen additional comparisons are made among the modern human population samples. Statistically significant differences are found for 16 of the 18 Neandertal vs. modern human comparisons but for only two of the 18 modern human comparisons. Statistical analyses repeated for subsamples of less worn teeth show a similar pattern. Because surface curvature is thought to affect perikymata spacing, we also conducted measurements to assess surface curvature in thirty teeth. Our analysis shows that surface curvature is not a factor in this lateral enamel growth difference between Neandertals and modern humans.     

The place of Neandertals in the evolution of hominid patterns of growth and development

This study uses the two developmental fields of dental maturation and femoral growth to determine if the pattern of growth and development in Neandertals (archaic Homo sapiens) was intermediate between that of Homo erectus and recent modern humans. Specimens used in the analysis included Neandertals and Upper Palaeolithic early modern Homo sapiens from Europe and individuals from two recent modern human populations. Ontogenetic data for the H. erectus adolescent KNM-WT 15000 and for Gorilla gorilla were included for comparison. Previous reports have indicated that H. erectus demonstrates a pattern of ontogeny characterized by earlier and more rapid linear growth than in modern humans. Results reported here demonstrate that Upper Paleolithic early modern Homo sapiens display a growth trajectory indistinguishable from that of recent modern humans. The pattern of Neandertal ontogeny is not intermediate between the pattern displayed in H. erectus and the derived pattern seen in the modern reference samples and the early modern H. sapiens sample. The Neandertal growth trajectory is consistent with either slow linear growth or advanced dental development.     

Is Central Asia the eastern outpost of the Neandertal range? A reassessment of the Teshik‐Tash child

Since its discovery in southeastern Uzbekistan in 1938, the Teshik-Tash child has been considered a Neandertal. Its affinity is important to studies of Late Pleistocene hominin growth and development as well as interpretations of the Central Asian Middle Paleolithic and the geographic distribution of Neandertals. A close examination of the original Russian monograph reveals the incompleteness of key morphologies associated with the cranial base and face and problems with the reconstruction of the Teshik-Tash cranium, making its Neandertal attribution less certain than previously assumed. This study reassesses the Neandertal status of Teshik-Tash 1 by comparing it to a sample of Neandertal, Middle and Upper Paleolithic modern humans, and recent human sub-adults. Separate examinations of the cranium and mandible are conducted using multinomial logistic regression and discriminant function analysis to assess group membership. Results of the cranial analysis group Teshik-Tash with Upper Paleolithic modern humans when variables are not size-standardized, while results of the mandibular analysis place the specimen with recent modern humans for both raw and size-standardized data. Although these results are influenced by limitations related to the incomplete nature of the comparative sample, they suggest that the morphology of Teshik-Tash 1 as expressed in craniometrics is equivocal. Although, further quantitative studies as well as additional sub-adult fossil finds from this region are needed to ascertain the morphological pattern of this specimen specifically, and Central Asian Middle Paleolithic hominins in general, these results challenge current characterizations of this territory as the eastern boundary of the Neandertal range during the Late Pleistocene.     

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.     

New discoveries and interpretations of hominid fossils and artifacts from Vindija Cave, Croatia

Beginning with excavations during the 1970s, Vindija Cave (Croatia) has yielded significant Middle and Upper Paleolithic fossil and archaeological finds. We report on seven recently identified hominid fossils, a newly associated partial hominid cranial vault from level G(3), nine possible bone retouchers, and a revised interpretation of the Mousterian artifact assemblage from the site. This new information reinforces our knowledge of the complex biocultural phenomena revealed in unit G and earlier deposits at Vindija. Six of the new hominid fossils derive from stratigraphic units G and I, while one lacks exact provenience. All specimens preserving diagnostic anatomy are from Neandertals. One of the postcranial remains, a radius fragment which exhibits Neandertal-like anatomy, comes from level G(1)and is congruent with the previously established association of Neandertals with an early Upper Paleolithic industry at the site. The partial cranial vault represents the most complete Neandertal from Vindija. The possible retouchers derive from unit G. Our analysis of these artifacts suggests that both percussion and pressure techniques may have been used by Neandertals in the final stage of tool production (retouching). This paper also presents a revision of the artifact analysis for late Mousterian level G(3). We separated raw materials into two main groups due to the differing ways that the materials fracture and the differing morphology of the debitage. The use of raw material in level G(3)is different from earlier Middle Paleolithic levels at Vindija. This indicates that the G(3)late Neandertals were making choices regarding source material somewhat more like the Upper Paleolithic people at the site. When interpreted within a larger regional framework, the Vindija archaeological and hominid fossil remains demonstrate a complex, mosaic pattern of biocultural change in the Late Pleistocene of south-central Europe.     

Pego do Diabo (Loures,Portugal): Dating the Emergence of Anatomical Modernity in Westernmost Eurasia

Background

Neandertals and the Middle Paleolithic persisted in the Iberian Peninsula south of the Ebro drainage system for several millennia beyond their assimilation/replacement elsewhere in Europe. As only modern humans are associated with the later stages of the Aurignacian, the duration of this persistence pattern can be assessed via the dating of diagnostic occurrences of such stages.

Methodology/Principal Findings

Using AMS radiocarbon and advanced pretreatment techniques, we dated a set of stratigraphically associated faunal samples from an Aurignacian III–IV context excavated at the Portuguese cave site of Pego do Diabo. Our results establish a secure terminus ante quem of ca.34,500 calendar years ago for the assimilation/replacement process in westernmost Eurasia. Combined with the chronology of the regional Late Mousterian and with less precise dating evidence for the Aurignacian II, they place the denouement of that process in the 37th millennium before present.

Conclusions/Significance

These findings have implications for the understanding of the emergence of anatomical modernity in the Old World as a whole, support explanations of the archaic features of the Lagar Velho child''s anatomy that invoke evolutionarily significant Neandertal/modern admixture at the time of contact, and counter suggestions that Neandertals could have survived in southwest Iberia until as late as the Last Glacial Maximum.     

Human remains from Valdegoba Cave (Huérmeces, Burgos, Spain).

Systematic excavations, begun in 1987, at the Valdegoba cave site in northern Spain have yielded the remains of five individuals associated with a Middle Paleolithic stone tool technology and Pleistocene fauna. A fragmentary mandible of an adolescent (VB1), preserving nearly a full set of teeth, exhibits a symphyseal tubercle and slight incurvatio mandibulae anterior on the external symphysis. Both the superior and inferior transverse tori are present on the internal aspect. A second individual (VB2) is represented by a set of ten deciduous teeth consistent with an age at death of 6-9 months. A proximal manual phalanx (VB3) displays a relatively broad head, a characteristic which is found in both Neandertals, as well as European Middle Pleistocene hominids. VB4 is a fourth metatarsal that lacks the distal epiphysis, indicating it comes from an adolescent individual, and has a relatively high robusticity index. Finally, VB5 is a fifth metatarsal of an adult. The VB1 mandible shows a combination of archaic characteristics as well as more specific Neandertal morphological traits. The VB2 deciduous teeth are very small, and both the metrics and morphology seem more consistent with a modern human classification. The postcranial elements are undiagnostic, U-Th dating has provided an age of >350 ka for the base of the sequence and a date of <73.2+/-5 ka for level 7, near the top. Faunal analysis and radiometric dates from other nearby Mousterian sites suggests that the Valdegoba site is correlative with oxygen isotope stages 3-6 on the Iberian peninsula, and an Upper Pleistocene age for the Valdegoba hominids seems most reasonable.     

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.