Thermoluminescence dating of the late Neanderthal remains from Saint-Césaire

Anatomically modern humans have long been thought to have been responsible for the Aurignacian and Chatelperronian industries of the early Upper Palaeolithic of Western Europe, whereas the Middle Palaeolithic Mousterian industry has been attributed to Neanderthals. The presence of both Middle and Upper Palaeolithic strata at Saint-Césaire in France offers an excellent opportunity for studying the cultural transition between the two. Saint-Césaire is the only Chatelperronian site that has yielded really diagnostic hominid fossils, and the discovery there of Neanderthal remains alongside Chatelperronian tools cast doubt on the exclusive association between industries and taxon. We report thermoluminescence dates for 20 burnt flints from the site. Those found near the Neanderthal remains were dated at 36,300 +/- 2,700 years BP (before present), making this specimen the youngest Neanderthal dated so far. This date places the stratum close in age to several French but much younger than some Spanish Aurignacian sites believed to have been occupied by modern humans. The possibility of contact between the West European Neanderthals and the intrusive modern humans who replaced them cannot therefore be excluded.     

Molecular analysis of Neanderthal DNA from the northern Caucasus

The expansion of premodern humans into western and eastern Europe approximately 40,000 years before the present led to the eventual replacement of the Neanderthals by modern humans approximately 28,000 years ago. Here we report the second mitochondrial DNA (mtDNA) analysis of a Neanderthal, and the first such analysis on clearly dated Neanderthal remains. The specimen is from one of the eastern-most Neanderthal populations, recovered from Mezmaiskaya Cave in the northern Caucasus. Radiocarbon dating estimated the specimen to be approximately 29,000 years old and therefore from one of the latest living Neanderthals. The sequence shows 3.48% divergence from the Feldhofer Neanderthal. Phylogenetic analysis places the two Neanderthals from the Caucasus and western Germany together in a clade that is distinct from modern humans, suggesting that their mtDNA types have not contributed to the modern human mtDNA pool. Comparison with modern populations provides no evidence for the multiregional hypothesis of modern human evolution.     

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.     

Radiocarbon dating of interstratified Neanderthal and early modern human occupations at the Chatelperronian type-site

The question of the coexistence and potential interaction between the last Neanderthal and the earliest intrusive populations of anatomically modern humans in Europe has recently emerged as a topic of lively debate in the archaeological and anthropological literature. Here we report the results of radiocarbon accelerator dating for what has been reported as an interstratified sequence of late Neanderthal and early anatomically modern occupations at the French type-site of the Chatelperronian, the Grotte des Fées de Chatelperron, in east-central France. The radiocarbon measurements seem to provide the earliest secure dates for the presence of Aurignacian technology--and from this, we infer the presence of anatomically modern human populations--in France.     

Analysis of one million base pairs of Neanderthal DNA

Neanderthals are the extinct hominid group most closely related to contemporary humans, so their genome offers a unique opportunity to identify genetic changes specific to anatomically fully modern humans. We have identified a 38,000-year-old Neanderthal fossil that is exceptionally free of contamination from modern human DNA. Direct high-throughput sequencing of a DNA extract from this fossil has thus far yielded over one million base pairs of hominoid nuclear DNA sequences. Comparison with the human and chimpanzee genomes reveals that modern human and Neanderthal DNA sequences diverged on average about 500,000 years ago. Existing technology and fossil resources are now sufficient to initiate a Neanderthal genome-sequencing effort.     

ESR dates for the hominid burial site of Es Skhul in Israel

The Middle East has been critical to our understanding of recent human evolution ever since the recovery of Neanderthal and early anatomically modern fossils from the caves of Tabun and Skhul (Mount Carmel) over 50 years ago. It was generally believed, on archaeological and morphological grounds, that middle eastern Neanderthals (such as those from Tabun, Amud and Kebara) probably dated from more than 50,000 years ago, whereas the earliest anatomically modern specimens (from Skhul and Qafzeh) probably dated from about 40,000 years. Recent thermoluminescence and electron spin resonance (ESR) determinations, however, have supported biostratigraphy in dating the Qafzeh deposits to an earlier part of the late Pleistocene, probably more than 90,000 years ago. These dates have been questioned on unspecified technical grounds, and it has also been argued that they create explanatory problems by separating the morphologically similar Qafzeh and Skhul samples by some 50,000 years, thus implying a long-term coexistence of early modern humans and Neanderthals in the area. Here we report the first radiometric dating analysis for Skhul, using ESR on bovine teeth from the hominid burial levels. Early uptake and linear uptake ages average 81 +/- 15 and 101 +/- 12 kyr respectively. These analyses suggest that the Skhul and Qafzeh samples are of a similar age and therefore it is possible that the presence of early modern humans in the area was episodic, rather than long-term during the early late Pleistocene.     

Late survival of Neanderthals at the southernmost extreme of Europe

The late survival of archaic hominin populations and their long contemporaneity with modern humans is now clear for southeast Asia. In Europe the extinction of the Neanderthals, firmly associated with Mousterian technology, has received much attention, and evidence of their survival after 35 kyr bp has recently been put in doubt. Here we present data, based on a high-resolution record of human occupation from Gorham's Cave, Gibraltar, that establish the survival of a population of Neanderthals to 28 kyr bp. These Neanderthals survived in the southernmost point of Europe, within a particular physiographic context, and are the last currently recorded anywhere. Our results show that the Neanderthals survived in isolated refuges well after the arrival of modern humans in Europe.     

Pleistocene Homo sapiens from Middle Awash,Ethiopia

The origin of anatomically modern Homo sapiens and the fate of Neanderthals have been fundamental questions in human evolutionary studies for over a century. A key barrier to the resolution of these questions has been the lack of substantial and accurately dated African hominid fossils from between 100,000 and 300,000 years ago. Here we describe fossilized hominid crania from Herto, Middle Awash, Ethiopia, that fill this gap and provide crucial evidence on the location, timing and contextual circumstances of the emergence of Homo sapiens. Radioisotopically dated to between 160,000 and 154,000 years ago, these new fossils predate classic Neanderthals and lack their derived features. The Herto hominids are morphologically and chronologically intermediate between archaic African fossils and later anatomically modern Late Pleistocene humans. They therefore represent the probable immediate ancestors of anatomically modern humans. Their anatomy and antiquity constitute strong evidence of modern-human emergence in Africa.     

A lost Neanderthal neonate found

Fossil remains of adult Neanderthals are well documented, but juvenile specimens are rare and information about them is scant. Here we identify a beautifully preserved skeleton that has been lost to science for almost 90 years as the Neanderthal neonate known as 'Le Moustier 2', which was originally found at Le Moustier in the Dordogne, southwest France. This find will be a rich source of data for studying the evolution of human ontogeny as well as the phylogenetic relationship between these extinct hominids and anatomically modern humans.     

Neanderthal cranial ontogeny and its implications for late hominid diversity

Homo neanderthalensis has a unique combination of craniofacial features that are distinct from fossil and extant 'anatomically modern' Homo sapiens (modern humans). Morphological evidence, direct isotopic dates and fossil mitochondrial DNA from three Neanderthals indicate that the Neanderthals were a separate evolutionary lineage for at least 500,000 yr. However, it is unknown when and how Neanderthal craniofacial autapomorphies (unique, derived characters) emerged during ontogeny. Here we use computerized fossil reconstruction and geometric morphometrics to show that characteristic differences in cranial and mandibular shape between Neanderthals and modern humans arose very early during development, possibly prenatally, and were maintained throughout postnatal ontogeny. Postnatal differences in cranial ontogeny between the two taxa are characterized primarily by heterochronic modifications of a common spatial pattern of development. Evidence for early ontogenetic divergence together with evolutionary stasis of taxon-specific patterns of ontogeny is consistent with separation of Neanderthals and modern humans at the species level.     

Human presence in the European Arctic nearly 40,000 years ago.

The transition from the Middle to the Upper Palaeolithic, approximately 40,000-35,000 radiocarbon years ago, marks a turning point in the history of human evolution in Europe. Many changes in the archaeological and fossil record at this time have been associated with the appearance of anatomically modern humans. Before this transition, the Neanderthals roamed the continent, but their remains have not been found in the northernmost part of Eurasia. It is generally believed that this vast region was not colonized by humans until the final stage of the last Ice Age some 13,000-14,000 years ago. Here we report the discovery of traces of human occupation nearly 40,000 years old at Mamontovaya Kurya, a Palaeolithic site situated in the European part of the Russian Arctic. At this site we have uncovered stone artefacts, animal bones and a mammoth tusk with human-made marks from strata covered by thick Quaternary deposits. This is the oldest documented evidence for human presence at this high latitude; it implies that either the Neanderthals expanded much further north than previously thought or that modern humans were present in the Arctic only a few thousand years after their first appearance in Europe.     

Unexpectedly recent dates for human remains from Vogelherd

The human skeletal remains from the Vogelherd cave in the Swabian Jura of southwestern Germany are at present seen as the best evidence that modern humans produced the artefacts of the early Aurignacian. Radiocarbon measurements from all the key fossils from Vogelherd show that these human remains actually date to the late Neolithic, between 3,900 and 5,000 radiocarbon years before present (bp). Although many questions remain unresolved, these results weaken the arguments for the Danube Corridor hypothesis--that there was an early migration of modern humans into the Upper Danube drainage--and strengthen the view that Neanderthals may have contributed significantly to the development of Upper Palaeolithic cultural traits independent of the arrival of modern humans.     

How Neanderthal molar teeth grew

Growth and development are both fundamental components of demographic structure and life history strategy. Together with information about developmental timing they ultimately contribute to a better understanding of Neanderthal extinction. Primate molar tooth development tracks the pace of life history evolution most closely, and tooth histology reveals a record of birth as well as the timing of crown and root growth. High-resolution micro-computed tomography now allows us to image complex structures and uncover subtle differences in adult tooth morphology that are determined early in embryonic development. Here we show that the timing of molar crown and root completion in Neanderthals matches those known for modern humans but that a more complex enamel-dentine junction morphology and a late peak in root extension rate sets them apart. Previous predictions about Neanderthal growth, based only on anterior tooth surfaces, were necessarily speculative. These data are the first on internal molar microstructure; they firmly place key Neanderthal life history variables within those known for modern humans.     

Direct dating of Early Upper Palaeolithic human remains from Mladec

The human fossil assemblage from the Mladec Caves in Moravia (Czech Republic) has been considered to derive from a middle or later phase of the Central European Aurignacian period on the basis of archaeological remains (a few stone artefacts and organic items such as bone points, awls, perforated teeth), despite questions of association between the human fossils and the archaeological materials and concerning the chronological implications of the limited archaeological remains. The morphological variability in the human assemblage, the presence of apparently archaic features in some specimens, and the assumed early date of the remains have made this fossil assemblage pivotal in assessments of modern human emergence within Europe. We present here the first successful direct accelerator mass spectrometry radiocarbon dating of five representative human fossils from the site. We selected sample materials from teeth and from one bone for 14C dating. The four tooth samples yielded uncalibrated ages of approximately 31,000 14C years before present, and the bone sample (an ulna) provided an uncertain more-recent age. These data are sufficient to confirm that the Mladec human assemblage is the oldest cranial, dental and postcranial assemblage of early modern humans in Europe and is therefore central to discussions of modern human emergence in the northwestern Old World and the fate of the Neanderthals.     

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.     

Neanderthals and the modern human colonization of Europe

The fate of the Neanderthal populations of Europe and western Asia has gripped the popular and scientific imaginations for the past century. Following at least 200,000 years of successful adaptation to the glacial climates of northwestern Eurasia, they disappeared abruptly between 30,000 and 40,000 years ago, to be replaced by populations all but identical to modern humans. Recent research suggests that the roots of this dramatic population replacement can be traced far back to events on another continent, with the appearance of distinctively modern human remains and artefacts in eastern and southern Africa.     

Huanglong Cave, a new late Pleistocene hominid site in Hubei Province, China

For the past two decades, the modern human origins debate has received significant interest from both the scientific community and the public. The two hypothe- ses “Out of Africa” and “Mutiregional evolution” are focuses of this debate[1-3]. In partic…     

Genetic history of an archaic hominin group from Denisova Cave in Siberia

Using DNA extracted from a finger bone found in Denisova Cave in southern Siberia, we have sequenced the genome of an archaic hominin to about 1.9-fold coverage. This individual is from a group that shares a common origin with Neanderthals. This population was not involved in the putative gene flow from Neanderthals into Eurasians; however, the data suggest that it contributed 4-6% of its genetic material to the genomes of present-day Melanesians. We designate this hominin population 'Denisovans' and suggest that it may have been widespread in Asia during the Late Pleistocene epoch. A tooth found in Denisova Cave carries a mitochondrial genome highly similar to that of the finger bone. This tooth shares no derived morphological features with Neanderthals or modern humans, further indicating that Denisovans have an evolutionary history distinct from Neanderthals and modern humans.     

Surprisingly rapid growth in Neanderthals

Life-history traits correlate closely with dental growth, so differences in dental growth within Homo can enable us to determine how somatic development has evolved and to identify developmental shifts that warrant species-level distinctions. Dental growth can be determined from the speed of enamel formation (or extension rate). We analysed the enamel extension rate in Homo antecessor (8 teeth analysed), Homo heidelbergensis (106), Homo neanderthalensis ('Neanderthals'; 146) and Upper Palaeolithic-Mesolithic Homo sapiens (100). Here we report that Upper Palaeolithic-Mesolithic H. sapiens shared an identical dental development pattern with modern humans, but that H. antecessor and H. heidelbergensis had shorter periods of dental growth. Surprisingly, Neanderthals were characterized by having the shortest period of dental growth. Because dental growth is an excellent indicator of somatic development, our results suggest that Neanderthals developed faster even than their immediate ancestor, H. heidelbergensis. Dental growth became longer and brain size increased from the Plio-Pleistocene in hominid evolution. Neanderthals, despite having a large brain, were characterized by a short period of development. This autapomorphy in growth is an evolutionary reversal, and points strongly to a specific distinction between H. sapiens and H. neanderthalensis.     

The effect of ancient population bottlenecks on human phenotypic variation

The origin and patterns of dispersal of anatomically modern humans are the focus of considerable debate. Global genetic analyses have argued for one single origin, placed somewhere in Africa. This scenario implies a rapid expansion, with a series of bottlenecks of small amplitude, which would have led to the observed smooth loss of genetic diversity with increasing distance from Africa. Analyses of cranial data, on the other hand, have given mixed results, and have been argued to support multiple origins of modern humans. Using a large data set of skull measurements and an analytical framework equivalent to that used for genetic data, we show that the loss in genetic diversity has been mirrored by a loss in phenotypic variability. We find evidence for an African origin, placed somewhere in the central/southern part of the continent, which harbours the highest intra-population diversity in phenotypic measurements. We failed to find evidence for a second origin, and we confirm these results on a large genetic data set. Distance from Africa accounts for an average 19-25% of heritable variation in craniometric measurements-a remarkably strong effect for phenotypic measurements known to be under selection.