European Neanderthals Research Articles

Long genetic and social isolation in Neanderthals before their extinction

Neanderthal genomes have been recovered from sites across Eurasia, painting an increasingly complex picture of their populations' structure that mostly indicates that late European Neanderthals belonged to a single metapopulation with no significant evidence of population structure. Here, we report the discovery of a late Neanderthal individual, nicknamed "Thorin," from Grotte Mandrin in Mediterranean France, and his genome. These dentognathic fossils, including a rare example of distomolars, are associated with a rich archeological record of Neanderthal final technological traditions in this region ∼50-42 thousand years ago. Thorin's genome reveals a relatively early divergence of ∼105 ka with other late Neanderthals. Thorin belonged to a population with a small group size that showed no genetic introgression with other known late European Neanderthals, revealing some 50 ka of genetic isolation of his lineage despite them living in neighboring regions. These results have important implications for resolving competing hypotheses about causes of the disappearance of the Neanderthals.

The east of the Neanderthal universe: a comparative analysis of Sukhaya Mechetka and Altai Micoquian complexes

Late European Neanderthals, associated with the Micoquian techno-complex, spread over vast area from Western Europe to the Altai. There were no sites found in the territories to the east of the Volga region. The key site of the Eastern Micoquian, Sukhaya Mechetka, which was discovered in 1951 in the Volgograd region, was for a long time the eastern outpost of Neanderthals in Eurasia. In the last five years, the results of paleogenetic analysis have confirmed the archaeological hypothesis of the migration of late European Neanderthals, the producers of the Micoquian, from Central and Eastern Europe to Altai. Until then, the genesis of Altai industries was associated with a wide range of European and Near Eastern techno-complexes, and anthropological remains were interpreted more in a Near Eastern context. In this regard, the industry of the Sukhaya Mechetka site, located on the route of the supposed migration of the Micoquians, acquires additional relevance in the research of the variability of the easternmost complexes of this cultural tradition. The attributive and technological analysis of tool kit from Sukhaya Mechetka was carried out and, using mathematical statistics methods, their results were compared with the available data on the Altai sites: Chagyrskaya and Okladnikov Caves. The new data were supplemented by the published materials from the Sukhaya Mechetka site. As a result, high level of technical-typological similarity of the compared industries in the context of primary flaking, bifacial tool production, tool typology and morphology was revealed. The structure of the Sukhaya Mechetka tool kit is closest to the Chagirskaya Сave, which is characterized by a large number of simple side-scrapers. At the same time, the site lacks of high-quality raw materials used to make bifacial items and intensively worked unifacial tools. In the Okladnikov Cave complexes this deficit is less. In composition and morphology of unifacial and bifacial tools, Sukhaya Mechetka is closest to the assemblages of the Antonovka I site and Chagyrkaya Cave. They share similarity with Okladnikov Cave in terms of a significant number of leaf-shaped and crescent-shaped tools. Some new characteristics were additionally determined for the assemblage of Sukhaya Mechetka. It was established that the primary decortications of cores was carried out outside the site. We also reconstructed the stages of production, rejuvenating and reshaping of bifaces on the site.

Identifying Palaeolithic birch tar production techniques: challenges from an experimental biomolecular approach

The intentional production of birch bark tar by European Neanderthals as early as 190,000 years ago plays an important role in discussions about the technological and behavioural complexity of Pleistocene hominins. However, research is hampered because it is currently unknown how Neanderthals were producing birch tar. There are several different techniques that could have been employed, but these differ in their apparent production complexity, time and resource efficiency. Identifying production processes in the archaeological record is therefore paramount for furthering research on the technical behavioural repertoire. Organic biomarkers, identified with Gas Chromatograph–Mass Spectrometry (GC–MS), have been used to identify possible production processes during the Neolithic. Here we test whether these biomarkers can also distinguish Palaeolithic (aceramic) tar production methods. We produced tar using five different methods and analysed their biomolecular composition with GC–MS. Our results show that the biomarkers used to distinguish Neolithic tar production strategies using ceramic technology cannot be reliably used to identify tar production processes using aceramic Palaeolithic techniques. More experimentation is required to produce a larger reference library of different tars for future comparisons. To achieve this, complete GC–MS datasets must also be made publicly available, as we have done with our data.

Neanderthal child's occipital from Baume Moula‐Guercy (Soyons, Ardèche, France)

AbstractObjectivesThis article provides an ontogenetically based comparative description of two immature occipital fragments from Baume Moula‐Guercy (MIS 5e) and examines their affinities to European and Middle Eastern Middle‐to‐Late Pleistocene (≈MIS 14–MIS ≈ 1) Homo.Materials and methodsDescription of the M‐S‐41 and M‐S‐61 occipital fragments (≈6–8 years) is based on observations of original fossils, casts, CT scans, literature descriptions, and virtual ectocranial and endocranial reconstructions. Our ontogenetically based sample represents a Preneanderthal‐Neanderthal group and a Homo sapiens group. These groups are subdivided into (1) Preneanderthals (≈MIS 14–9), Early Neanderthals (MIS 7–5e), and Late Neanderthals (MIS 5d‐3), and (2) Middle (MIS 5), Upper (MIS 3–1), and Late (MIS ≈ 1) Paleolithic H. sapiens. Measurements and developmental age determinations follow standard techniques.ResultsBased on the M‐S‐41/M‐S‐61 composite, the strongly convex upper occipital scale flattens into the vertical suprainiac fossa, as in immature Early Neanderthals. A doubled suprainiac fossa expressing a weakly developed bounding torus and pocking of the central region is typical of immature Neanderthals. The transverse torus is thickened medially, lacks significant lateral development, has a laterally placed protuberance, and is concave medially. A linear (triangular) tubercle, as opposed to an external occipital protuberance, is present. An occipital bun is absent.DiscussionThe occipital remains from Moula‐Guercy make a substantial contribution to the record of Neanderthal cranial evolution because immature Early Neanderthal occipitals are rare, fragmentary, and likely represent only two paleodemes (Krapina, La Chaise‐de‐Vouthon). The Moula‐Guercy occipital possesses characteristics well‐established in European Neanderthals by MIS 7, and it is most similar to the occipitals of other Early Neanderthals.

Недеструктивная объемная микроскопия в изучении особенностей биологической адаптации среди ископаемых и современных Homo

We apply non-destructive volumetric technique for evaluation of vascular net density in compact bone. Segmentation of Haversian canals and accurate estimation of their relative volume fraction gives the opportunity to estimate the density and capacity of the vascular system of the bone tissue in a proxy way. Radiological images of dorsal compact of medial and distal phalanges were studied (Neanderthals from Altai and Europe; CroMagnons; recent humans of Arctic and African origin). Results. High vascularization of compact clearly demonstrates that European Neanderthals and the majority of AMH were adapted to the Glacial Age. But Kostenki 14 (Eastern Europe) and Strashnaya 4 (Altai) have features in common with humans of recent tropic origin. Specimens of Neanderthals which lived in Altai region where glaciers never occurred are more diverse. Hypothetically the milder climate favored the existence of a wider adaptive norm reflecting patterns of former environments. Somewhat ambiguous morphological patterns of Denisova 9 and Okladnikov 2, 5 may reflect various episodes of Neanderthal migration to Siberia and their hybridization with people of tropical ancestry, most probably the early AMH. Chagyrskaya Neanderthal demonstrates a hyper-cold adaptation. Conclusion. Diversity in microstructural patterns of the Pleistocene people’s skeletal system gives evidence that both Neanderthals and CroMagnons were polymorphic in their adaptive reactions and presumably included both warm and cold adapted forms.

A Neanderthal from the Central Western Zagros, Iran. Structural reassessment of the Wezmeh 1 maxillary premolar

Wezmeh Cave, in the Kermanshah region of Central Western Zagros, Iran, produced a Late Pleistocene faunal assemblage rich in carnivorans along with a human right maxillary premolar, Wezmeh 1, an unerupted tooth from an 8 ± 2 year-old individual. Uranium-series analyses of the fauna by alpha spectrometry provided age estimates between 70 and 11 ka. Crown dimensions place the tooth specimen at the upper limits of Late Pleistocene human ranges of variation. Wezmeh 1 metameric position (most likely a P3) remains uncertain and only its surficial morphology has been described so far. Accordingly, we used microfocus X-ray tomography (12.5 μm isotropic voxel size) to reassess the metameric position and taxonomic attribution of this specimen. We investigated its endostructural features and quantified crown tissue proportions. Topographic maps of enamel thickness (ET) distribution were also generated, and semilandmark-based geometric morphometric analyses of the enamel-dentine junction (EDJ) were performed. We compared Wezmeh 1 with unworn/slightly-moderately worn P3 and P4 of European Neanderthals, Middle Paleolithic modern humans from Qafzeh, an Upper Paleolithic premolar, and Holocene humans. The results confirm that Wezmeh 1 represents a P3. Based on its internal conformation and especially EDJ shape, Wezmeh 1 aligns closely with Neanderthals and is distinct from the fossil and extant modern human pattern of our comparative samples. Wezmeh 1 is thus the first direct evidence of Neanderthal presence on the western margin of the Iranian Plateau.

Metric and morphological comparison between the Arago (France) and Atapuerca-Sima de los Huesos (Spain) dental samples, and the origin of Neanderthals

The variability observed in the growing Middle Pleistocene hominin fossil record of Europe continues to trigger much debate on taxonomic issues and the biological processes that gave rise to Neanderthals. Here we present a metric and morphological comparative study of the dental samples recovered from the sites of Arago (southeast France) and Sima de los Huesos (SH) in the Sierra de Atapuerca (northern Spain). These sites are key to providing answers to these questions since they have yielded the largest hominin samples so far recovered for this time period. Despite the geographical proximity of the two sites and the contemporaneity of their hominin assemblages, we have observed remarkable metric and morphological differences between the teeth at Arago and SH. Whereas the SH teeth present an almost morphological identity with European Neanderthals, the Arago teeth exhibit a combination of plesiomorphic as well as some Neanderthal-derived features. In addition, the Arago crown dimensions are remarkably larger than those from SH, the differences being statistically significant for most variables. We hypothesize that during the Middle Pleistocene the European continent was settled at different points in time by hominin groups coming from Southwest Asia, probably from a common mother population evolving in this latter region. These first settlers can be identified by their more plesiomorphic morphology, whereas the most recent settlers are closer in appearance to Neanderthals. In addition, genetic processes such as isolation, genetic drift, directional adaptation or hybridization would have given rise to the puzzle we observe in the current fossil record.

Human Colonization of Asia in the Late Pleistocene

Human Colonization of Asia in the Late Pleistocene

Drawings of Representational Images by Upper Paleolithic Humans and their Absence in Neanderthals Reflect Historical Differences in Hunting Wary Game

AbstractOne characteristic of the transition from the Middle Paleolithic to the Upper Paleolithic in Europe was the emergence of representational charcoal drawings and engravings by Aurignacian and Gravettian artists. European Neanderthals never engaged in representational drawing during the Middle and Early Upper Paleolithic, a property that might reflect less developed visuomotor coordination. This article postulates a causal relationship between an evolved ability of anatomically modern humans to throw spears accurately while hunting and their ability to draw representational images from working memory. Unlike Neanderthals, archaic and anatomically modern humans in sub-Saharan Africa adopted longer-range hunting practices using hand-cast spears as a compensation for the emergence of increasingly wary game. For Neanderthals, paleoclimatic fluctuations likely precluded consistent hunting of cold-adapted game, a property making game more approachable for close-range hunting with thrusting spears. As evidence of less historical wariness of humans, many of the species hunted by Neanderthals were eventually domesticated. Due to strong sources of natural selection on archaic and anatomically modern humans for effective hunting, the parietal cortex that integrates visual imagery and motor coordination expanded progressively, yielding the globular shape of the human cranium that is not evident in Neanderthals. To characterize how the cognitive properties employed for throwing spears and drawing line work are similar, the Upper Paleolithic drawings of animals in Chauvet cave, France, are discussed in the speculative context of how these artists engaged simultaneously in overt attention to guide their hand movements and covert attention to their mental images during the drawing process.

How Did Hominins Adapt to Ice Age Europe without Fire?

Analyses of archaeological material recovered from several Middle Paleolithic sites in southwest France have provided strong corroborating data on Neanderthal use of fire. Both direct and indirect data show that Neanderthals in this region were frequently and/or intensively using fire during warmer periods, but such evidence declines significantly in occupations that took place during colder periods. One possible explanation for this pattern is that it reflects the inability of Western European Neanderthals to make fire, simply because natural sources of fire occur much more frequently during warmer climatic periods. Regardless of the explanation, the long periods of diminished evidence of fire shows that, unlike modern humans, these hominins were not obligate fire users, and this fact in itself raises important questions of how they adapted, physiologically and/or technologically, to the generally harsh glacial conditions of the middle latitude of Europe and to reduced energy returns typical of raw food....

Robusticity of Hand Phalanges: Relevance to the Origin of the Altai Neanderthals

Cross-sectional geometry of middle phalanges of hand digits 2–4 in fi ve European and Asian Neanderthals (La Ferrassie 1, Kiik-Koba 1, Okladnikov 2 and 5, and Chagyrskaya 16-3-12) and fi ve Cro-Magnons (Kostenki 14, Telmanovskaya TII 175 and TII 173, Sungir 1, and Abri Pataud 26227) was estimated by means of microtomography. Both Neanderthals and Cro-Magnons reveal a wide range of individual variability in the inner robusticity indices. Both the most robust and the most gracile variants in Neanderthals were recorded in the Altai (Okladnikov and Chagyrskaya caves, respectively), confi rming previous observations about the high morphological diversity among Neanderthals in that area and the presence of at least two morphological variants among them. In European Upper Paleolithic Homo sapiens, inner phalangeal robusticity is generally higher than in Neanderthals, attaining medullary stenosis in the Kostenki 14 male. Neither sex nor age nor even mechanical stress appear to have affected robusticity. Hyperrobust variants were recognized in both Cro-Magnons and Neanderthals of supposedly hybrid origin. Genetic studies suggest that Kostenki 14 belonged to an ancestral European metapopulation, which had absorbed some Neanderthal admixture. The ancestors of the Altai Neanderthals, on the other hand, included not only Denisovans but also early anatomically modern humans before their migration to Siberia. Extreme phalangeal robusticity in Middle and Upper Paleolithic Eurasians, then, might be a legacy of early anatomically modern humans.

Ancient gene flow from early modern humans into Eastern Neanderthals.

It has been shown that Neanderthals contributed genetically to modern humans outside Africa 47,000-65,000 years ago. Here we analyse the genomes of a Neanderthal and a Denisovan from the Altai Mountains in Siberia together with the sequences of chromosome 21 of two Neanderthals from Spain and Croatia. We find that a population that diverged early from other modern humans in Africa contributed genetically to the ancestors of Neanderthals from the Altai Mountains roughly 100,000 years ago. By contrast, we do not detect such a genetic contribution in the Denisovan or the two European Neanderthals. We conclude that in addition to later interbreeding events, the ancestors of Neanderthals from the Altai Mountains and early modern humans met and interbred, possibly in the Near East, many thousands of years earlier than previously thought.

Levantine cranium from Manot Cave (Israel) foreshadows the first European modern humans.

A key event in human evolution is the expansion of modern humans of African origin across Eurasia between 60 and 40 thousand years (kyr) before present (bp), replacing all other forms of hominins. Owing to the scarcity of human fossils from this period, these ancestors of all present-day non-African modern populations remain largely enigmatic. Here we describe a partial calvaria, recently discovered at Manot Cave (Western Galilee, Israel) and dated to 54.7±5.5kyr bp (arithmetic mean±2 standard deviations) by uranium-thorium dating, that sheds light on this crucial event. The overall shape and discrete morphological features of the Manot 1 calvaria demonstrate that this partial skull is unequivocally modern. It is similar in shape to recent African skulls as well as to European skulls from the Upper Palaeolithic period, but different from most other early anatomically modern humans in the Levant. This suggests that the Manot people could be closely related to the first modern humans who later successfully colonized Europe. Thus, the anatomical features used to support the 'assimilation model' in Europe might not have been inherited from European Neanderthals, but rather from earlier Levantine populations. Moreover, at present, Manot 1 is the only modern human specimen to provide evidence that during the Middle to Upper Palaeolithic interface, both modern humans and Neanderthals contemporaneously inhabited the southern Levant, close in time to the likely interbreeding event with Neanderthals.

Neanderthal hand and foot remains from Moula‐Guercy, Ardèche, France

The hand and foot remains from Moula-Guercy cave (Ardèche, France) comprise 24 specimens of Eemian age (ca. 120 ka). The specimens include primarily complete elements, which are rare among the Moula-Guercy postcrania. The hand remains have several characteristic Neanderthal traits including a laterally facing (parasagittally oriented) second metacarpal-capitate articulation, a short styloid process, a wide proximal articular surface on the third metacarpal, and absolutely expanded apical tuberosities on the distal hand phalanges relative to modern humans. The foot remains include several incomplete elements along with an antimeric pair of naviculars, a medial cuneiform and cuboid, and a single complete element from each of the distal segments (one each: metatarsal, proximal foot phalanx, intermediate foot phalanx, distal foot phalanx). Consistent among the specimens are relatively wide diaphyses for length in the metatarsals and phalanges and large and prominent muscle attachments, both consistent with previously published Neanderthal morphology. The hand and foot collection from Moula-Guercy is an important dataset for future studies of Neanderthal functional morphology, dexterity, and behavior as it represents a previously undersampled time period for European Neanderthals.

Neanderthal Graphic Behavior

This paper presents a study of a pecked pebble from the Middle Paleolithic recovered more than 30 years ago in Axlor rockshelter in the Spanish Basque Country. At the time of the discovery, the piece was described as being deliberately modified, but since then it has been either ignored or described only as problematic evidence of Neanderthal symbolic behavior. In this work, we provide a new description and additional documentation of the piece, and we discuss its anthropic nature and the possibility that its meaning can be related to the small but credible record of graphic behavior known for early hominin groups prior to the Upper Paleolithic in Europe. The linear design theme of the Axlor piece is similar to that of other items made by European Neanderthals. These artifacts confirm the capability of pre-Homo sapiens sapiens humans to create and use portable “art.” This tradition is evidence of the emergence of “behavioral modernity” among late Neanderthals.

Mitochondrial DNA polymorphisms shared between modern humans and neanderthals: Adaptive convergence or evidence for interspecific hybridization?

An analysis of the variability of the nucleotide sequences in the mitochondrial genome of modern humans, neanderthals, Denisovans, and other primates has shown that there are shared polymorphisms at positions 2758 and 7146 between modern Homo sapiens (in phylogenetic cluster L2'3'4'5'6) and Homo neanderthalensis (in the group of European neanderthals younger than 48000 years). It is suggested that the convergence may be due to adaptive changes in the mitochondrial genomes of modern humans and neanderthals or interspecific hybridization associated with mtDNA recombination.

The Neanderthals of Okladnikov Cave Altai: Environment and Diet Based on Isotopic Analysis*

The discovery of a Mousteroid industry in the northwestern Altai represents a major finding in Eurasian Paleolithic studies. Isotopic analyses of bone collagen of late Western European Neanderthals have repeatedly indicated that they were extremely carnivorous. However, no corresponding information was previously available for the Neanderthals of the Altai. With this in mind, an isotopic study was conducted to assess the diet of the Middle Paleolithic inhabitants of Okladnikov Cave. Collagen was extracted from seven well preserved bone samples of adults and children and 13С/12С and 15N/14N ratios were calculated. The δ13С content suggests that all individuals were members of a local trophic chain based on plants of the moderate climatic zone. However, the δ15N content demonstrates that their trophic level was high. These findings suggest that the lifestyle of Neanderthals in the northwestern Altai was similar to that of the late European Neanderthals. Both evidently subsisted by hunting large herbivores.

A multidisciplinary approach to reconstructing the chronology and environment of southwestern European Neanderthals: the contribution of Teixoneres cave (Moià, Barcelona, Spain)

According to pollen analysis and the 18O-isotope curve, the first part of the Late Pleistocene (ca 128–30 ka) is mainly characterized by a dynamic that alternates cold phases (Heinrich Events) and temperate phases (interstadials). These rapid fluctuations provide the context for the Neanderthal occupations in the northeastern part of the Iberian Peninsula. In this paper we present the chronological, environmental and climatic data obtained by analyzing the pollen, the charcoal, the small vertebrates (amphibians, squamates and small mammals) and the large-mammal dental wear at the Neanderthal site of Teixoneres cave, Northeastern Iberia. Levels II and III from this cavity have provided Mousterian industries and other evidence of Neanderthal occupations, such as cut-marks in large-mammal bones. A multiproxy study such as this constitutes a new approach to the chronological, environmental and climatic context in which Neanderthal populations lived in southwestern Europe (Iberian Peninsula). The results allow us to establish a relative chronology for these two levels of between ca 30–90 ka and show that they are associated with different environmental and climatic conditions: temperate and humid for Level III and cold and dry for Level II. This demonstrates that the Neanderthals were well adapted to the territory that they occupied, irrespective of the climatic conditions.

European Neanderthal stone hunting weapons reveal complex behaviour long before the appearance of modern humans

Stone weapon points constitute a major innovation appearing at the end of the Middle Pleistocene in Europe, Africa and the Near East; that is, among both sapiens and Neanderthal populations. The microscopic analysis of the stone weapons used by Neanderthal groups in Atlantic southern Europe suggests that this technology was widespread and became a recurrent behaviour within organised strategies developed by these societies. In this southern region of Europe, stone weapon hunting technology appears at an early time (about 150 ka, OIS 6) and is associated with the hunting of large mammals. This behaviour can be recognised in a geomorphologically complex region and at a time of great environmental change (OIS 6–5–4). The fact that these innovations were used by European Neanderthals long before the spread of anatomically modern humans in the area is of great evolutionary significance.

The earliest evidence for anatomically modern humans in northwestern Europe

The earliest anatomically modern humans in Europe are thought to have appeared around 43,000-42,000 calendar years before present (43-42 kyr cal BP), by association with Aurignacian sites and lithic assemblages assumed to have been made by modern humans rather than by Neanderthals. However, the actual physical evidence for modern humans is extremely rare, and direct dates reach no farther back than about 41-39 kyr cal BP, leaving a gap. Here we show, using stratigraphic, chronological and archaeological data, that a fragment of human maxilla from the Kent's Cavern site, UK, dates to the earlier period. The maxilla (KC4), which was excavated in 1927, was initially diagnosed as Upper Palaeolithic modern human. In 1989, it was directly radiocarbon dated by accelerator mass spectrometry to 36.4-34.7 kyr cal BP. Using a Bayesian analysis of new ultrafiltered bone collagen dates in an ordered stratigraphic sequence at the site, we show that this date is a considerable underestimate. Instead, KC4 dates to 44.2-41.5 kyr cal BP. This makes it older than any other equivalently dated modern human specimen and directly contemporary with the latest European Neanderthals, thus making its taxonomic attribution crucial. We also show that in 13 dental traits KC4 possesses modern human rather than Neanderthal characteristics; three other traits show Neanderthal affinities and a further seven are ambiguous. KC4 therefore represents the oldest known anatomically modern human fossil in northwestern Europe, fills a key gap between the earliest dated Aurignacian remains and the earliest human skeletal remains, and demonstrates the wide and rapid dispersal of early modern humans across Europe more than 40 kyr ago.