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.     

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.     

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.     

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.     

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.     

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.     

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.     

DNA sequences from the quagga, an extinct member of the horse family

To determine whether DNA survives and can be recovered from the remains of extinct creatures, we have examined dried muscle from a museum specimen of the quagga, a zebra-like species (Equus quagga) that became extinct in 1883 (ref. 1). We report that DNA was extracted from this tissue in amounts approaching 1% of that expected from fresh muscle, and that the DNA was of relatively low molecular weight. Among the many clones obtained from the quagga DNA, two containing pieces of mitochondrial DNA (mtDNA) were sequenced. These sequences, comprising 229 nucleotide pairs, differ by 12 base substitutions from the corresponding sequences of mtDNA from a mountain zebra, an extant member of the genus Equus. The number, nature and locations of the substitutions imply that there has been little or no postmortem modification of the quagga DNA sequences, and that the two species had a common ancestor 3-4 Myr ago, consistent with fossil evidence concerning the age of the genus Equus.     

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 new species of great ape from the late Miocene epoch in Ethiopia

With the discovery of Ardipithecus, Orrorin and Sahelanthropus, our knowledge of hominid evolution before the emergence of Pliocene species of Australopithecus has significantly increased, extending the hominid fossil record back to at least 6 million years (Myr) ago. However, because of the dearth of fossil hominoid remains in sub-Saharan Africa spanning the period 12-7 Myr ago, nothing is known of the actual timing and mode of divergence of the African ape and hominid lineages. Most genomic-based studies suggest a late divergence date-5-6 Myr ago and 6-8 Myr ago for the human-chimp and human-gorilla splits, respectively-and some palaeontological and molecular analyses hypothesize a Eurasian origin of the African ape and hominid clade. We report here the discovery and recognition of a new species of great ape, Chororapithecus abyssinicus, from the 10-10.5-Myr-old deposits of the Chorora Formation at the southern margin of the Afar rift. To the best of our knowledge, these are the first fossils of a large-bodied Miocene ape from the African continent north of Kenya. They exhibit a gorilla-sized dentition that combines distinct shearing crests with thick enamel on its 'functional' side cusps. Visualization of the enamel-dentine junction by micro-computed tomography reveals shearing crest features that partly resemble the modern gorilla condition. These features represent genetically based structural modifications probably associated with an initial adaptation to a comparatively fibrous diet. The relatively flat cuspal enamel-dentine junction and thick enamel, however, suggest a concurrent adaptation to hard and/or abrasive food items. The combined evidence suggests that Chororapithecus may be a basal member of the gorilla clade, and that the latter exhibited some amount of adaptive and phyletic diversity at around 10-11 Myr ago.     

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.     

Evolution and environment in the Hominoidea.

Between 10 and 20 million years ago, a variety of hominoid primates lived in Africa, Europe and Asia. The question of which of these, if any, lie closest to the ancestries of humans and modern apes remains a lively source of debate. Recent fossil discoveries, though, shed light on the environments in which the various groups of hominoid emerged and, it is hoped, on their evolution. But the lack of a hominid fossil record before about 5 million years ago--and any fossil record for the African apes--is still a frustrating barrier.     

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.     

A Middle Palaeolithic human hyoid bone

The origin of human language, and in particular the question of whether or not Neanderthal man was capable of language/speech, is of major interest to anthropologists but remains an area of great controversy. Despite palaeoneurological evidence to the contrary, many researchers hold to the view that Neanderthals were incapable of language/speech, basing their arguments largely on studies of laryngeal/basicranial morphology. Studies, however, have been hampered by the absence of unambiguous fossil evidence. We now report the discovery of a well-preserved human hyoid bone from Middle Palaeolithic layers of Kebara Cave, Mount Carmel, Israel, dating from about 60,000 years BP. The bone is almost identical in size and shape to the hyoid of present-day populations, suggesting that there has been little or no change in the visceral skeleton (including the hyoid, middle ear ossicles, and inferentially the larynx) during the past 60,000 years of human evolution. We conclude that the morphological basis for human speech capability appears to have been fully developed during the Middle Palaeolithic.     

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.     

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.     

内蒙古砧子山墓地古人的线粒体DNA多态性分析

为了研究内蒙古元上都遗址砧子山墓地古代人群的遗传结构及其可能来源,对该墓地古人的DNA进行了抽提、扩增和测序,获得了10个个体的线粒体DNA高可变一区序列.结合现代东亚、北亚、中亚和欧洲人的线粒体DNA数据进行了系统发育分析和多维尺度分析.研究结果表明:埋藏在砧子山墓地的元代居民为汉族人,主要是来自中国北方地区的汉族.本研究为揭示元代的复杂社会结构和人群历史动态提供了新的方法.     

Woody cover and hominin environments in the past 6 million years

The role of African savannahs in the evolution of early hominins has been debated for nearly a century. Resolution of this issue has been hindered by difficulty in quantifying the fraction of woody cover in the fossil record. Here we show that the fraction of woody cover in tropical ecosystems can be quantified using stable carbon isotopes in soils. Furthermore, we use fossil soils from hominin sites in the Awash and Omo-Turkana basins in eastern Africa to reconstruct the fraction of woody cover since the Late Miocene epoch (about 7 million years ago). (13)C/(12)C ratio data from 1,300 palaeosols at or adjacent to hominin sites dating to at least 6 million years ago show that woody cover was predominantly less than ～40% at most sites. These data point to the prevalence of open environments at the majority of hominin fossil sites in eastern Africa over the past 6 million years.