Homo sapiens reached the higher latitudes of Europe by 45,000 years ago.

The Middle to Upper Palaeolithic transition in Europe is associated with the regional disappearance of Neanderthals and the spread of Homo sapiens. Late Neanderthals persisted in western Europe several millennia after the occurrence of H. sapiens in eastern Europe1. Local hybridization between the two groups occurred2, but not on all occasions3. Archaeological evidence also indicates the presence of several technocomplexes during this transition, complicating our understanding and the association of behavioural adaptations with specific hominin groups4. One such technocomplex for which the makers are unknown is the Lincombian-Ranisian-Jerzmanowician (LRJ), which has been described in northwestern and central Europe5-8. Here we present the morphological and proteomic taxonomic identification, mitochondrial DNA analysis and direct radiocarbon dating of human remains directly associated with an LRJ assemblage at the site Ilsenhöhle in Ranis (Germany). These human remains are among the earliest directly dated Upper Palaeolithic H. sapiens remains in Eurasia. We show that early H. sapiens associated with the LRJ were present in central and northwestern Europe long before the extinction of late Neanderthals in southwestern Europe. Our results strengthen the notion of a patchwork of distinct human populations and technocomplexes present in Europe during this transitional period.

Ancient human DNA recovered from a Palaeolithic pendant.

Artefacts made from stones, bones and teeth are fundamental to our understanding of human subsistence strategies, behaviour and culture in the Pleistocene. Although these resources are plentiful, it is impossible to associate artefacts to specific human individuals1 who can be morphologically or genetically characterized, unless they are found within burials, which are rare in this time period. Thus, our ability to discern the societal roles of Pleistocene individuals based on their biological sex or genetic ancestry is limited2-5. Here we report the development of a non-destructive method for the gradual release of DNA trapped in ancient bone and tooth artefacts. Application of the method to an Upper Palaeolithic deer tooth pendant from Denisova Cave, Russia, resulted in the recovery of ancient human and deer mitochondrial genomes, which allowed us to estimate the age of the pendant at approximately 19,000-25,000 years. Nuclear DNA analysis identifies the presumed maker or wearer of the pendant as a female individual with strong genetic affinities to a group of Ancient North Eurasian individuals who lived around the same time but were previously found only further east in Siberia. Our work redefines how cultural and genetic records can be linked in prehistoric archaeology.

Initial Upper Palaeolithic humans in Europe had recent Neanderthal ancestry.

Modern humans appeared in Europe by at least 45,000 years ago1-5, but the extent of their interactions with Neanderthals, who disappeared by about 40,000 years ago6, and their relationship to the broader expansion of modern humans outside Africa are poorly understood. Here we present genome-wide data from three individuals dated to between 45,930 and 42,580 years ago from Bacho Kiro Cave, Bulgaria1,2. They are the earliest Late Pleistocene modern humans known to have been recovered in Europe so far, and were found in association with an Initial Upper Palaeolithic artefact assemblage. Unlike two previously studied individuals of similar ages from Romania7 and Siberia8 who did not contribute detectably to later populations, these individuals are more closely related to present-day and ancient populations in East Asia and the Americas than to later west Eurasian populations. This indicates that they belonged to a modern human migration into Europe that was not previously known from the genetic record, and provides evidence that there was at least some continuity between the earliest modern humans in Europe and later people in Eurasia. Moreover, we find that all three individuals had Neanderthal ancestors a few generations back in their family history, confirming that the first European modern humans mixed with Neanderthals and suggesting that such mixing could have been common.

Pluridisciplinary evidence for burial for the La Ferrassie 8 Neandertal child.

The origin of funerary practices has important implications for the emergence of so-called modern cognitive capacities and behaviour. We provide new multidisciplinary information on the archaeological context of the La Ferrassie 8 Neandertal skeleton (grand abri of La Ferrassie, Dordogne, France), including geochronological data -14C and OSL-, ZooMS and ancient DNA data, geological and stratigraphic information from the surrounding context, complete taphonomic study of the skeleton and associated remains, spatial information from the 1968-1973 excavations, and new (2014) fieldwork data. Our results show that a pit was dug in a sterile sediment layer and the corpse of a two-year-old child was laid there. A hominin bone from this context, identified through Zooarchaeology by Mass Spectrometry (ZooMS) and associated with Neandertal based on its mitochondrial DNA, yielded a direct 14C age of 41.7-40.8 ka cal BP (95%), younger than the 14C dates of the overlying archaeopaleontological layers and the OSL age of the surrounding sediment. This age makes the bone one of the most recent directly dated Neandertals. It is consistent with the age range for the Châtelperronian in the site and in this region and represents the third association of Neandertal taxa to Initial Upper Palaeolithic lithic technocomplex in Western Europe. A detailed multidisciplinary approach, as presented here, is essential to advance understanding of Neandertal behavior, including funerary practices.

A wolf from Gravettian site Pavlov I, Czech Republic: Approach to skull pathology.

OBJECTIVE: Describe pathological features on internal and external aspects of the skull of an ancient grey wolf. MATERIALS: Wolf remains that were found at the southwestern settlement Area A of Gravettian site Pavlov I. METHODS: Visual observation and description; microcomputed tomography; porosity and fragmentation indices for internal and external skull features; histological section of the fourth upper premolar tooth. RESULTS: Dorsally, the sagittal crest revealed bone healing and remodeling. The sagittal lesion differential diagnosis was blunt trauma with or without fracture. Ventrally, otic region pathology included severe proliferation and lysis (osteomyelitis). The pathology was not resolvable among differential (microbial) causes of osteomyelitis, although other potential etiologies were ruled out. CONCLUSIONS: Probable first report of otic region osteomyelitis in an ancient grey wolf. SIGNIFICANCE: The proximity of the wolf remains to human-related findings, and presence of red ochre and shells, suggest human involvement in the burial. LIMITATIONS: This is a single specimen with differential diagnoses that were not resolvable to a single definitive diagnosis. SUGGESTIONS FOR FURTHER RESEARCH: Further investigation of the possible anthropological significance of the burial circumstances.

Initial Upper Palaeolithic Homo sapiens from Bacho Kiro Cave, Bulgaria.

The Middle to Upper Palaeolithic transition in Europe witnessed the replacement and partial absorption of local Neanderthal populations by Homo sapiens populations of African origin1. However, this process probably varied across regions and its details remain largely unknown. In particular, the duration of chronological overlap between the two groups is much debated, as are the implications of this overlap for the nature of the biological and cultural interactions between Neanderthals and H. sapiens. Here we report the discovery and direct dating of human remains found in association with Initial Upper Palaeolithic artefacts2, from excavations at Bacho Kiro Cave (Bulgaria). Morphological analysis of a tooth and mitochondrial DNA from several hominin bone fragments, identified through proteomic screening, assign these finds to H. sapiens and link the expansion of Initial Upper Palaeolithic technologies with the spread of H. sapiens into the mid-latitudes of Eurasia before 45 thousand years ago3. The excavations yielded a wealth of bone artefacts, including pendants manufactured from cave bear teeth that are reminiscent of those later produced by the last Neanderthals of western Europe4-6. These finds are consistent with models based on the arrival of multiple waves of H. sapiens into Europe coming into contact with declining Neanderthal populations7,8.

The age of the hominin fossils from Jebel Irhoud, Morocco, and the origins of the Middle Stone Age.

The timing and location of the emergence of our species and of associated behavioural changes are crucial for our understanding of human evolution. The earliest fossil attributed to a modern form of Homo sapiens comes from eastern Africa and is approximately 195 thousand years old, therefore the emergence of modern human biology is commonly placed at around 200 thousand years ago. The earliest Middle Stone Age assemblages come from eastern and southern Africa but date much earlier. Here we report the ages, determined by thermoluminescence dating, of fire-heated flint artefacts obtained from new excavations at the Middle Stone Age site of Jebel Irhoud, Morocco, which are directly associated with newly discovered remains of H. sapiens. A weighted average age places these Middle Stone Age artefacts and fossils at 315 ± 34 thousand years ago. Support is obtained through the recalculated uranium series with electron spin resonance date of 286 ± 32 thousand years ago for a tooth from the Irhoud 3 hominin mandible. These ages are also consistent with the faunal and microfaunal assemblages and almost double the previous age estimates for the lower part of the deposits. The north African site of Jebel Irhoud contains one of the earliest directly dated Middle Stone Age assemblages, and its associated human remains are the oldest reported for H. sapiens. The emergence of our species and of the Middle Stone Age appear to be close in time, and these data suggest a larger scale, potentially pan-African, origin for both.

On the local Mousterian origin of the Châtelperronian: Integrating typo-technological, chronostratigraphic and contextual data.

Across Europe the period 45-40 ka (thousands of years ago) is associated with several technological changes, including the emergence of the Châtelperronian technocomplex in France and northern Spain. The Châtelperronian, stratigraphically located between the Mousterian and Aurignacian, is characterized by Upper Palaeolithic features, such as volumetric blade reduction, curved backed blades, end-scrapers, bladelets, bone tools and ornaments. Concurrently, repeated, though debated, associations with Neanderthal remains and Mousterian elements suggest a local technological development. Following recent critiques and cumulating technological studies, this paper provides data-driven contextualisations of the Châtelperronian and late Mousterian archaeological records and a primary comparative assessment of a major linking element, backed knives, to re-assess the origin of the Châtelperronian. The results demonstrate the challenging nature of the 50-35 ka record, with many interpretive problems caused by poorly recorded excavations, resulting in only 25 well-contextualised assemblages from the claimed 143 Châtelperronian find spots. These 25 assemblages facilitate more detailed chronostratigraphic and typo-technological assessments and show that the Châtelperronian has a homogenous set of technologies and tools. A similar evaluation of the late Mousterian indicates a wide-ranging late Neanderthal skill set, commonly including laminar blank production and backing. Further, conceptual similarities were noted both in blank selection and edge modification between Mousterian and Châtelperronian backed knives, alongside their near-absence in other, contemporaneous technocomplexes. A Europe-wide contextualisation shows that while the current coarse-grained record still allows for several potential scenarios, the data throughout this paper point towards a most parsimonious model of a Châtelperronian made by Neanderthals, with roots in the late Middle Palaeolithic technological skill set. However, this change seems triggered by early arrivals of modern humans either indirectly, through stimulus diffusion, or directly, after ca. 42 ka. Fully testing this model requires an ongoing focus on site formation and assemblage integrity, alongside in-depth analyses of recently excavated assemblages and existing collections.

New chronology for Ksâr 'Akil (Lebanon) supports Levantine route of modern human dispersal into Europe.

Modern human dispersal into Europe is thought to have occurred with the start of the Upper Paleolithic around 50,000-40,000 y ago. The Levantine corridor hypothesis suggests that modern humans from Africa spread into Europe via the Levant. Ksâr 'Akil (Lebanon), with its deeply stratified Initial (IUP) and Early (EUP) Upper Paleolithic sequence containing modern human remains, has played an important part in the debate. The latest chronology for the site, based on AMS radiocarbon dates of shell ornaments, suggests that the appearance of the Levantine IUP is later than the start of the first Upper Paleolithic in Europe, thus questioning the Levantine corridor hypothesis. Here we report a series of AMS radiocarbon dates on the marine gastropod Phorcus turbinatus associated with modern human remains and IUP and EUP stone tools from Ksâr 'Akil. Our results, supported by an evaluation of individual sample integrity, place the EUP layer containing the skeleton known as "Egbert" between 43,200 and 42,900 cal B.P. and the IUP-associated modern human maxilla known as "Ethelruda" before ∼ 45,900 cal B.P. This chronology is in line with those of other Levantine IUP and EUP sites and demonstrates that the presence of modern humans associated with Upper Paleolithic toolkits in the Levant predates all modern human fossils from Europe. The age of the IUP-associated Ethelruda fossil is significant for the spread of modern humans carrying the IUP into Europe and suggests a rapid initial colonization of Europe by our species.

Rapid dental development in a Middle Paleolithic Belgian Neanderthal.

The evolution of life history (pace of growth and reproduction) was crucial to ancient hominin adaptations. The study of dental development facilitates assessment of growth and development in fossil hominins with greater precision than other skeletal analyses. During tooth formation, biological rhythms manifest in enamel and dentine, creating a permanent record of growth rate and duration. Quantification of these internal and external incremental features yields developmental benchmarks, including ages at crown completion, tooth eruption, and root completion. Molar eruption is correlated with other aspects of life history. Recent evidence for developmental differences between modern humans and Neanderthals remains ambiguous. By measuring tooth formation in the entire dentition of a juvenile Neanderthal from Scladina, Belgium, we show that most teeth formed over a shorter time than in modern humans and that dental initiation and eruption were relatively advanced. By registering manifestations of stress across the dentition, we are able to present a precise chronology of Neanderthal dental development that differs from modern humans. At 8 years of age at death, this juvenile displays a degree of development comparable with modern human children who are several years older. We suggest that age at death in juvenile Neanderthals should not be assessed by comparison with modern human standards, particularly those derived from populations of European origin. Moreover, evidence from the Scladina juvenile and other similarly aged hominins suggests that a prolonged childhood and slow life history are unique to Homo sapiens.

Neanderthals in central Asia and Siberia.

Morphological traits typical of Neanderthals began to appear in European hominids at least 400,000 years ago and about 150,000 years ago in western Asia. After their initial appearance, such traits increased in frequency and the extent to which they are expressed until they disappeared shortly after 30,000 years ago. However, because most fossil hominid remains are fragmentary, it can be difficult or impossible to determine unambiguously whether a fossil is of Neanderthal origin. This limits the ability to determine when and where Neanderthals lived. To determine how far to the east Neanderthals ranged, we determined mitochondrial DNA (mtDNA) sequences from hominid remains found in Uzbekistan and in the Altai region of southern Siberia. Here we show that the DNA sequences from these fossils fall within the European Neanderthal mtDNA variation. Thus, the geographic range of Neanderthals is likely to have extended at least 2,000 km further to the east than commonly assumed.

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 Châtelperronian 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, Châtelperronian, Aurignacian, and Gravettian layers. Previously, a child's temporal bone from the Châtelperronian Layer Xb was recognized as belonging to a Neandertal; however, most of the teeth from Châtelperronian layers VIII-X remain unpublished. We describe the dental remains from the Châtelperronian 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 Châtelperronian 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 Châtelperronian industry at Arcy-sur-Cure.