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.     

Early dispersal of modern humans in Europe and implications for Neanderthal behaviour

The appearance of anatomically modern humans in Europe and the nature of the transition from the Middle to Upper Palaeolithic are matters of intense debate. Most researchers accept that before the arrival of anatomically modern humans, Neanderthals had adopted several 'transitional' technocomplexes. Two of these, the Uluzzian of southern Europe and the Chatelperronian of western Europe, are key to current interpretations regarding the timing of arrival of anatomically modern humans in the region and their potential interaction with Neanderthal populations. They are also central to current debates regarding the cognitive abilities of Neanderthals and the reasons behind their extinction. However, the actual fossil evidence associated with these assemblages is scant and fragmentary, and recent work has questioned the attribution of the Chatelperronian to Neanderthals on the basis of taphonomic mixing and lithic analysis. Here we reanalyse the deciduous molars from the Grotta del Cavallo (southern Italy), associated with the Uluzzian and originally classified as Neanderthal. Using two independent morphometric methods based on microtomographic data, we show that the Cavallo specimens can be attributed to anatomically modern humans. The secure context of the teeth provides crucial evidence that the makers of the Uluzzian technocomplex were therefore not Neanderthals. In addition, new chronometric data for the Uluzzian layers of Grotta del Cavallo obtained from associated shell beads and included within a Bayesian age model show that the teeth must date to ~45,000-43,000 calendar years before present. The Cavallo human remains are therefore the oldest known European anatomically modern humans, confirming a rapid dispersal of modern humans across the continent before the Aurignacian and the disappearance of Neanderthals.     

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.     

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.     

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.     

Early human occupation of the Red Sea coast of Eritrea during the last interglacial

The geographical origin of modern humans is the subject of ongoing scientific debate. The 'multiregional evolution' hypothesis argues that modern humans evolved semi-independently in Europe, Asia and Africa between 100,000 and 40,000 years ago, whereas the 'out of Africa' hypothesis contends that modern humans evolved in Africa between 200 and 100 kyr ago, migrating to Eurasia at some later time. Direct palaeontological, archaeological and biological evidence is necessary to resolve this debate. Here we report the discovery of early Middle Stone Age artefacts in an emerged reef terrace on the Red Sea coast of Eritrea, which we date to the last interglacial (about 125 kyr ago) using U-Th mass spectrometry techniques on fossil corals. The geological setting of these artefacts shows that early humans occupied coastal areas and exploited near-shore marine food resources in East Africa by this time. Together with similar, tentatively dated discoveries from South Africa this is the earliest well-dated evidence for human adaptation to a coastal marine environment, heralding an expansion in the range and complexity of human behaviour from one end of Africa to the other. This new, wide-spread adaptive strategy may, in part, signal the onset of modern human behaviour, which supports an African origin for modern humans by 125 kyr ago.     

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.     

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.     

The earliest record of human activity in northern Europe

The colonization of Eurasia by early humans is a key event after their spread out of Africa, but the nature, timing and ecological context of the earliest human occupation of northwest Europe is uncertain and has been the subject of intense debate. The southern Caucasus was occupied about 1.8 million years (Myr) ago, whereas human remains from Atapuerca-TD6, Spain (more than 780 kyr ago) and Ceprano, Italy (about 800 kyr ago) show that early Homo had dispersed to the Mediterranean hinterland before the Brunhes-Matuyama magnetic polarity reversal (780 kyr ago). Until now, the earliest uncontested artefacts from northern Europe were much younger, suggesting that humans were unable to colonize northern latitudes until about 500 kyr ago. Here we report flint artefacts from the Cromer Forest-bed Formation at Pakefield (52 degrees N), Suffolk, UK, from an interglacial sequence yielding a diverse range of plant and animal fossils. Event and lithostratigraphy, palaeomagnetism, amino acid geochronology and biostratigraphy indicate that the artefacts date to the early part of the Brunhes Chron (about 700 kyr ago) and thus represent the earliest unequivocal evidence for human presence north of the Alps.     

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.     

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.     

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.     

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 dating evidence for early modern humans at Border Cave in South Africa

The archaeological and hominid site of Border Cave (KwaZulu, South Africa) has a stratigraphic sequence covering the Middle and Later Stone Ages (MSA and LSA). It has been proposed that four hominid specimens recovered there (BC1 and BC2 of uncertain provenance, and BC3 and BC5 recovered from MSA layers) represent very early examples of anatomically modern humans, supporting an early late-Pleistocene appearance of modern Homo sapiens in Africa. This early appearance, however, has been questioned, largely because of doubts about the stratigraphic positions associated with the specimens and because of the lack of a reliable chronology for the stratigraphic sequence. We now report on the first comprehensive radiometric dating analysis of Border Cave, using electron spin resonance (ESR) on teeth within sediment layers. BC3 is likely to be approximately 70-80 kyr old, and BC5, 50-65 kyr old. BC1 and BC2 are almost certainly less than 90 kyr old. These results, although younger than some age estimates, support the early occurrence of anatomically modern humans at Border Cave. In addition, our results suggest that the Howiesons Poort lithic industry (approximately 45-75 kyr) and the MSA-LSA transition (approximately 35 kyr) are younger than often believed.     

A complete human pelvis from the Middle Pleistocene of Spain.

The Middle Pleistocene site of Sima de los Huesos in Sierra de Atapuerca, Spain, has yielded around 2,500 fossils from at least 33 different hominid individuals. These have been dated at more than 200,000 years ago and have been classified as ancestors of Neanderthals. An almost complete human male pelvis (labelled Pelvis 1) has been found, which we associate with two fragmentary femora. Pelvis 1 is robust and very broad with a very long superior pubic ramus, marked iliac flare, and a long femoral neck. This pattern is probably the primitive condition from which modern humans departed. A modern human newborn would pass through the birth canal of Pelvis 1 and this would be even larger in a female individual. We estimate the body mass of this individual at 95 kg or more. Using the cranial capacities of three specimens from Sima de los Huesos, the encephalization quotients are substantially smaller than in Neanderthals and modern humans.     

Early human use of marine resources and pigment in South Africa during the Middle Pleistocene

Genetic and anatomical evidence suggests that Homo sapiens arose in Africa between 200 and 100 thousand years (kyr) ago, and recent evidence indicates symbolic behaviour may have appeared approximately 135-75 kyr ago. From 195-130 kyr ago, the world was in a fluctuating but predominantly glacial stage (marine isotope stage MIS6); much of Africa was cooler and drier, and dated archaeological sites are rare. Here we show that by approximately 164 kyr ago (+/-12 kyr) at Pinnacle Point (on the south coast of South Africa) humans expanded their diet to include marine resources, perhaps as a response to these harsh environmental conditions. The earliest previous evidence for human use of marine resources and coastal habitats was dated to approximately 125 kyr ago. Coincident with this diet and habitat expansion is an early use and modification of pigment, probably for symbolic behaviour, as well as the production of bladelet stone tool technology, previously dated to post-70 kyr ago. Shellfish may have been crucial to the survival of these early humans as they expanded their home ranges to include coastlines and followed the shifting position of the coast when sea level fluctuated over the length of MIS6.     

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.     

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.     

Early stone technology on Flores and its implications for Homo floresiensis

In the Soa Basin of central Flores, eastern Indonesia, stratified archaeological sites, including Mata Menge, Boa Lesa and Kobatuwa (Fig. 1), contain stone artefacts associated with the fossilized remains of Stegodon florensis, Komodo dragon, rat and various other taxa. These sites have been dated to 840-700 kyr bp (thousand years before present). The authenticity of the Soa Basin artefacts and their provenance have been demonstrated by previous work, but to quell lingering doubts, here we describe the context, attributes and production modes of 507 artefacts excavated at Mata Menge. We also note specific similarities, and apparent technological continuity, between the Mata Menge stone artefacts and those excavated from Late Pleistocene levels at Liang Bua cave, 50 km to the west. The latter artefacts, dated to between 95-74 and 12 kyr ago, are associated with the remains of a dwarfed descendent of S. florensis, Komodo dragon, rat and a small-bodied hominin species, Homo floresiensis, which had a brain size of about 400 cubic centimetres. The Mata Menge evidence negates claims that stone artefacts associated with H. floresiensis are so complex that they must have been made by modern humans (Homo sapiens).     

Stratigraphic placement and age of modern humans from Kibish, Ethiopia

In 1967 the Kibish Formation in southern Ethiopia yielded hominid cranial remains identified as early anatomically modern humans, assigned to Homo sapiens. However, the provenance and age of the fossils have been much debated. Here we confirm that the Omo I and Omo II hominid fossils are from similar stratigraphic levels in Member I of the Kibish Formation, despite the view that Omo I is more modern in appearance than Omo II. 40Ar/39Ar ages on feldspar crystals from pumice clasts within a tuff in Member I below the hominid levels place an older limit of 198 +/- 14 kyr (weighted mean age 196 +/- 2 kyr) on the hominids. A younger age limit of 104 +/- 7 kyr is provided by feldspars from pumice clasts in a Member III tuff. Geological evidence indicates rapid deposition of each member of the Kibish Formation. Isotopic ages on the Kibish Formation correspond to ages of Mediterranean sapropels, which reflect increased flow of the Nile River, and necessarily increased flow of the Omo River. Thus the 40Ar/39Ar age measurements, together with the sapropel correlations, indicate that the hominid fossils have an age close to the older limit. Our preferred estimate of the age of the Kibish hominids is 195 +/- 5 kyr, making them the earliest well-dated anatomically modern humans yet described.