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Direct radiocarbon dating and DNA analysis of the Darra-i-Kur (Afghanistan) human temporal bone

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Abstract

The temporal bone discovered in the 1960s from the Darra-i-Kur cave in Afghanistan is often cited as one of the very few Pleistocene human fossils from Central Asia. Here we report the first direct radiocarbon date for the specimen and the genetic analyses of DNA extracted and sequenced from two areas of the bone. The new radiocarbon determination places the find to ∼4500 cal BP (∼2500 BCE) contradicting an assumed Palaeolithic age of ∼30,000 years, as originally suggested. The DNA retrieved from the specimen originates from a male individual who carried mitochondrial DNA of the modern human type. The petrous part yielded more endogenous ancient DNA molecules than the squamous part of the same bone. Molecular dating of the Darra-i-Kur mitochondrial DNA sequence corroborates the radiocarbon date and suggests that the specimen is younger than previously thought. Taken together, the results consolidate the fact that the human bone is not associated with the Pleistocene-age deposits of Darra-i-Kur; instead it is intrusive, possibly re-deposited from upper levels dating to much later periods (Neolithic). Despite its Holocene age, the Darra-i-Kur specimen is, so far, the first and only ancient human from Afghanistan whose DNA has been sequenced.

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... For Darra-i-Kur, we analyzed a single-stranded DNA library (L5082) at the Max Planck Institute for Evolutionary Anthropology in Leipzig, Germany, generated as part of a previous study (78). The previous study only analyzed mitochondrial DNA, and for the current study, we enriched the library for sequences overlapping the same panel of about 1.2 million nuclear targets using two rounds of hybridization capture (7,18,19). ...
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Ancient human movements through Asia Ancient DNA has allowed us to begin tracing the history of human movements across the globe. Narasimhan et al. identify a complex pattern of human migrations and admixture events in South and Central Asia by performing genetic analysis of more than 500 people who lived over the past 8000 years (see the Perspective by Schaefer and Shapiro). They establish key phases in the population prehistory of Eurasia, including the spread of farming peoples from the Near East, with movements both westward and eastward. The people known as the Yamnaya in the Bronze Age also moved both westward and eastward from a focal area located north of the Black Sea. The overall patterns of genetic clines reflect similar and parallel patterns in South Asia and Europe. Science , this issue p. eaat7487 ; see also p. 981
... A temporal bone had been found in the excavation, morphologically attributed to AMH (Angel, 1972). A recent DNA sampling of the bone confirms that it belonged to a male AMH, but it was a Neolithic individual, and was associated with the Mousterian assemblage because of taphonomic activity (Douka et al., 2017). (Dupree and Davis, 1972). ...
Thesis
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The Altai range, in southern Russia, has yielded an important series of prehistoric assemblages in various contexts (caves, shelters and open-air sites). Recent anthropological and archaeological studies have established the significance of this area, with complex peopling events involving at least three different human species, Neanderthals, Modern Humans and Denisovans, the latter being exclusively associated with Altai assemblages. The cultural background of these hominins’ occupation is already well defined for the beginning of the Upper Palaeolithic. This study aims to better characterize the previous period’s productions, through one of its important cultural features, the Levallois technology.To address this issue, we have undertaken a review of material coming from the some of the key Altai sequences, while trying to reconstruct the different chaînes opératoires implemented for the production of the desired products that had been previously recognized as Levallois. The analysed artefacts cover a large time span, from Early Middle Palaeolithic (Denisova, stratum 22 of the Central Chamber, RTL dated to 220-280 ky) to layers associated with Upper Palaeolithic (Ust’-Kanskaya, strata 3 to 1), and come from both caves and open-air sites. This allowed us to establish a chronological comparison, as well as regional. Results have shown that the Levallois assemblages of the region are quite homogenous; also, that Levallois technology may not have been present in Altai as early as it has been previously claimed, with a difference of ~100.000 years; and finally, that it is mostly analogous to what we can find in neighbouring regions. These extra-regional common features probably express contacts and exchanges to and from the Altai region.
... This is clearly not the case, as has been shown previously for cranial morphologies (64). Similar limitations apply to inferring population affiliations of isolated fossil hominin labyrinths (65)(66)(67). One reason is that within-population variation is larger than between-population variation. ...
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The cavity system of the inner ear—the so-called bony labyrinth—houses the senses of balance and hearing. This structure is embedded in dense petrous bone, fully formed by birth and generally well preserved in human skeletal remains, thus providing a rich source of morphological information about past populations. Here we show that labyrinthine morphology tracks genetic distances and geography in an isolation-by-distance model with dispersal from Africa. Because petrous bones have become prime targets of ancient DNA recovery, we propose that all destructive studies first acquire high-resolution 3D computed-tomography data prior to any invasive sampling. Such data will constitute an important archive of morphological variation in past and present populations, and will permit individual-based genotype–phenotype comparisons.The dispersal of modern humans from Africa is now well documented with genetic data that track population history, as well as gene flow between populations. Phenetic skeletal dat
... This is clearly not the case, as has been shown previously for cranial morphologies (64). Similar limitations apply to inferring population affiliations of isolated fossil hominin labyrinths (65)(66)(67). One reason is that within-population variation is larger than between-population variation. ...
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Significance The cavity system of the inner ear—the so-called bony labyrinth—houses the senses of balance and hearing. This structure is embedded in dense petrous bone, fully formed by birth and generally well preserved in human skeletal remains, thus providing a rich source of morphological information about past populations. Here we show that labyrinthine morphology tracks genetic distances and geography in an isolation-by-distance model with dispersal from Africa. Because petrous bones have become prime targets of ancient DNA recovery, we propose that all destructive studies first acquire high-resolution 3D computed-tomography data prior to any invasive sampling. Such data will constitute an important archive of morphological variation in past and present populations, and will permit individual-based genotype–phenotype comparisons.
... The assessment of the state of conservation of bone may help in deciding about its strength for excavation or to preserve it for further studies, for palaeopathological investigations, to reconstruct burial episodes, funeral rituals, and post-mortem events which may have caused dispersion of the skeletal elements (Baron et al. 1996;Trueman and Martill 2002;Nielsen-Marsh et al. 2007;Smith et al. 2007;Turner-Walker 2008;Hollund et al. 2012;De Boer et al. 2013;Booth 2016). Moreover, it may indicate alterations of the isotopic and chemical signature of bone, crucial for palaeodietary and palaeoclimatologic studies, to reconstruct the lifestyle of past populations and their mobility (DeNiro and Weiner 1988;Ambrose 1990; Lee-Thorp and van der Merwe 1991;Wright and Schwarcz 1996;Stott et al. 1999;van Klinken 1999;Hedges 2003;King et al. 2011;Assis et al. 2016;Douka et al. 2017). ...
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Diagenetic modifications in human bones from the early-medieval cemetery discovered in the garden of Vratislavs’ Palace, in the central Malá Strana district of Prague, have been investigated combining histological analysis and instrumental analysis with X-ray diffraction, infrared, and ³¹P NMR spectroscopy. A total of 15 ribs samples were collected for the study. One sample belonged to a child, whereas, of the other samples from adults, 7 belonged to males, 5 to females, and for 2 the sex attribution was uncertain. A diagenetic pathway common to most of the studied samples was considered the result of a burial environment characterized by a nearly static water regime, with limited temperature excursions, moderately oxic to suboxic, and with pH fluctuations around the limit of apatite recrystallization window, in agreement with the fine textured clay-rich soil, its low hydraulic conductivity, and the measured soil pH. A second pattern, related to variations in the microenvironment, interested a limited number of samples with poorer histological preservation. This was interpreted as the result of higher pH and a better oxygenated environment, which favoured mineral recrystallization. Further reactivation of deterioration processes probably occurred later in some of the graves perturbed by works conducted in the seventeenth century. This work highlights the complementarity of the information obtained from the adopted techniques in order to gain insights into the post-mortem fate of the human remains and their sedimentary environment. In this respect, the quantification of the amount of phosphorus in the amorphous hydrated layer of apatite provided a unique type of information on the mineral component of bone and its reorganization during diagenesis, revealing that a relevant fraction can survive diagenesis, at variance with what previously supposed.
... regions of the Subcontinent (Nishiaki & Akazawa 2018: 5) most probably because of the absence of research in the region. They are known from the cave of Darra-i-Kur in north-eastern Afghanistan (Dupree & Davis 1972), where they are still undated (see Douka et al. 2017), as well as in the easternmost part of South Khorasan (Iran) (Nasab 2011: fig. 1), not far from one of the suggested dispersal, migratory routes across the Iranian plateau towards Afghanistan (Nasab et al. 2013: fig. ...
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This paper discusses the importance of the discovery of one déjeté Levallois tool from the surface of a dark grey and black patinated gravel terrace located ca. 500 m southwest of the Neolithic site of Sheri Khan Tarakai in the Bannu Basin (Khyber Pakhtunkhwa, Pakistan), and provides a detailed geomorphological description of the area where it was found. The Neolithic site rests on a large gravelly fan, at present terraced and dismembered by small seasonal streams. Scatters of black varnished pebbles, at the top of a thick ochre silt of possible alluvial origin, cover its surface. Amongst the numerous siliceous gravels forming the deposit, some are of a good quality chert, whose source can be found in the Tertiary Sulaiman Formation. The typological characteristics of the tool, the chert employed for its manufacture, its location and the presence of black patina on its cortex are all important elements that contribute to the definition of the Pleistocene period during which pebble terraces formed. The tool comes from a region where Middle Palaeolithic artefacts had never been found before, though the re-analysis of old collections would suggest their presence as far as the course of the Indus in Lower Sindh. Moreover, its discovery contributes to the study of the southeastern spread of the Middle Palaeolithic Levallois technique, an important topic that still needs to be fully understood.
... The cochlea has since become a targeted skeletal element in human ancient DNA analyses 5,6,8,[23][24][25][31][32][33] . While the majority of recent ancient DNA studies use the petrous pyramid to provide the necessary bone powder for DNA extraction, they do not describe the specific process used to isolate and obtain bone fractions for paleogenomic analyses, and thus leave the reader uncertain as to whether the cochlea was in fact the part of the osseous inner ear used. ...
Article
The cortical bone that forms the structure of the cochlea, part of the osseous labyrinth of the inner ear, is now one of the most frequently used skeletal elements in analyses of human ancient DNA. However, there is currently no published, standardized method for its sampling. This protocol describes the preparation of bone powder from the cochlea of fragmented skulls in which the petrous pyramid of the temporal bone is accessible. Using a systematic process of bone removal based on distinct anatomical landmarks and the identification of relevant morphological features, a petrous pyramid is cleaned with a sandblaster, and the cochlea is located, isolated, and reduced to a homogeneous bone powder. All steps are carried out in dedicated ancient DNA facilities, thus reducing the introduction of contamination. This protocol requires an understanding of ancient DNA clean-room procedures and basic knowledge of petrous pyramid anatomy. In 50–65 min, it results in bone powder with endogenous DNA yields that can exceed those from teeth and other bones by up to two orders of magnitude. Compared with drilling methods, this method facilitates a more precise targeting of the cochlea, allows the user to visually inspect the cochlea and remove any residual sediment before the generation of bone powder, and confines the damage to the inner ear region and surface of the petrous portion of fragmentary crania. © 2019, The Author(s), under exclusive licence to Springer Nature Limited.
... This form of interpretation has already led to misunderstandings on numerous occasions. For instance, a bone from the Darra-i-Kur cave in Afghanistan, initially assumed to be from the Palaeolithic (30,000 BP) (Dupree 1972) and often cited as one of the very few Pleistocene human fossils from Central Asia, was recently radiocarbon-dated to the Neolithic (4,500 BP) (Douka et al. 2017). Though a major limitation of radiocarbon dating is the high amount of collagen extraction (500 mg), the authors could have employed TPS (Esposito et al. 2019) to support their dating, which, excepting Sample #656, were unjustified. ...
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Recently, Mikheyev et al. (2019) have produced a preprint study describing the genomes of nine Khazars archeologically dated from the 7th to the 9th centuries found in the Rostov county in modern-day Russia. Skull morphology indicated a mix of "Caucasoid" and "Mongoloid" shapes. The authors compared the samples to ancient and contemporary samples to study the genetic makeup of the Khazars and their genetic legacy and addressed the question of the relationships between the Khazar and Ashkenazic Jews. A careful examination reveals grave concerns regarding all the aspects of the study from the identification of the "Khazar" samples, the choice of environment for ancient DNA sequencing, and the analyses. The authors did not disclose the data used in their study, and their methodology is incoherent. We demonstrate that their analyses yield nonsensical results and argue that none of the claims made in this study are supported by the data unequivocally. Provided the destruction of the bone samples and the irreproducibility of the analyses, even by the forgivable standards of the field, this study is irreplicable, wasteful, and misleading.
... C dating), past lifeways (e.g., palaeodietary reconstruction, weaning practices), taxonomic identification, and occasionally for screening (e.g., amino acid racemization) (DeNiro and Weiner 1988;Ambrose 1991;Poinar et al. 1996;Collins et al. 1999Collins et al. , 2009van Klinken 1999;Asara et al. 2007;Buckley et al. 2009;Douka et al. 2017). ...
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This paper presents the characteristics of bone diagenesis in a secondary commingled Mycenaean burial in Kastrouli (Phocis, Greece) through the histological (light microscopy), physical (FTIR-ATR), and biochemical (collagen) analysis of seventeen human (including two petrous bones) and seven animal bones. Post-mortem modifications in bone microstructure, bioapatite, and collagen were characteristic of burial environments with seasonal groundwater and temperature fluctuations. The two human petrous bones displayed a lack of microscopic focal destruction (MFD) sites and a generally good histological preservation, but although a small sample size, did not show any better bioapatite and collagen preservation compared with human femora. Intra-site variations were defined by three main diagenetic patterns that display differences in histological modifications, crystallinity changes, and collagen degradation. These different patterns were either related to different microenvironment conditions and/or influenced by possible differences in the early taphonomic histories experienced by bones prior to secondary deposition. Further, this study highlights the importance of infrared splitting factor (IRSF), carbonate/phosphate ratio (C/P), and general histological index (GHI) for the qualitative assessment of archaeological bone, and the potential use of amide/phosphate ratio (Am/P) as a collagen predictor.
... Some laboratories have used microCT scanning, both to preserve data from petrous bone, and to guide their drilling to minimize destruction of the specimen 11 . Unfortunately, such methods have not been adopted as a standard, partly because individual groups tend to focus on their own research agenda rather than on the bigger picture. ...
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We present a high-quality genome sequence of a Neanderthal woman from Siberia. We show that her parents were related at the level of half-siblings and that mating among close relatives was common among her recent ancestors. We also sequenced the genome of a Neanderthal from the Caucasus to low coverage. An analysis of the relationships and population history of available archaic genomes and 25 present-day human genomes shows that several gene flow events occurred among Neanderthals, Denisovans and early modern humans, possibly including gene flow into Denisovans from an unknown archaic group. Thus, interbreeding, albeit of low magnitude, occurred among many hominin groups in the Late Pleistocene. In addition, the high-quality Neanderthal genome allows us to establish a definitive list of substitutions that became fixed in modern humans after their separation from the ancestors of Neanderthals and Denisovans.
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Excavations at the Chagyrskaya Cave site in Gorny Altai have revealed a Mousteroid industry along with fragmented human remains. This study focuses on a left ulna from stratum 6a. Its size, proportions, symptoms of disease, and indicators of muscular activity, point to Neanderthal af nities. The bone is large, linking the individual with certain Near Eastern Neanderthal males such as Shanidar. Symptoms of what might be diagnosed as Forestier disease suggest likewise. © 2013, Siberian Branch of Russian Academy of Sciences, Institute of Archaeology and Ethnography of the Siberian Branch of the Russian Academy of Sciences.
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Since its discovery in southeastern Uzbekistan in 1938, the Teshik-Tash child has been considered a Neandertal. Its affinity is important to studies of Late Pleistocene hominin growth and development as well as interpretations of the Central Asian Middle Paleolithic and the geographic distribution of Neandertals. A close examination of the original Russian monograph reveals the incompleteness of key morphologies associated with the cranial base and face and problems with the reconstruction of the Teshik-Tash cranium, making its Neandertal attribution less certain than previously assumed. This study reassesses the Neandertal status of Teshik-Tash 1 by comparing it to a sample of Neandertal, Middle and Upper Paleolithic modern humans, and recent human sub-adults. Separate examinations of the cranium and mandible are conducted using multinomial logistic regression and discriminant function analysis to assess group membership. Results of the cranial analysis group Teshik-Tash with Upper Paleolithic modern humans when variables are not size-standardized, while results of the mandibular analysis place the specimen with recent modern humans for both raw and size-standardized data. Although these results are influenced by limitations related to the incomplete nature of the comparative sample, they suggest that the morphology of Teshik-Tash 1 as expressed in craniometrics is equivocal. Although, further quantitative studies as well as additional sub-adult fossil finds from this region are needed to ascertain the morphological pattern of this specimen specifically, and Central Asian Middle Paleolithic hominins in general, these results challenge current characterizations of this territory as the eastern boundary of the Neandertal range during the Late Pleistocene.
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Two Middle Paleolithic cave sites in the Altai – Okladnikov and Chagyrskaya – have yielded dental remains (mostly isolated teeth) of individuals of various ages. A newly discovered mandibular fragment with teeth from Chagyrskaya Cave reveals a Neanderthal trait combination: anterior fossa and epicristid (midtrigonid crest) on molars, metaconid and crest on premolars. The totality of dental traits support the conclusion previously drawn on the basis of postcranial characters: Altai Neanderthals appear to be intermediate between other Eurasian Neanderthals and anatomically modern humans.
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To eliminate possible confusion in the reporting of isotopic abundances on noncorresponding scales, the Commission on Atomic Weights and Isotopic Abundances recommended at the 37th General Assembly at Lisbon, Portugal that (i) 2H/1H relative ratios of all substances be expressed relative to VSMOW (Vienna Standard Mean Ocean Water) on a scale such that 2H/1H of SLAP (Standard Light Antarctic Precipitation) is 0.572 times that of VSMOW, (ii) 13C/12C relative ratios of all substances be expressed relative to VPDB (Vienna Peedee belemnite) on a scale such that 13C/12C of NBS 19 carbonate is 1.00195 times that of VPDB, and (iii) 18O/16O ratios of all substances be expressed relative to either VSMOW or VPDB on scales such that 18O/16O of SLAP is 0.9445 times that of VSMOW.
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Tracing continuity of evolving humans with the aid of their scanty skeletal remains is an intricate puzzle. In the specific case of Indian sub-continent, the so-far oldest human ancestor, the Narmada hominid, has wider spatial distance both from the antecedents on one side and the descendents on the other side. Ramapithecus, a geographically closely located primate earlier considered of hominidae affinity, is now far out of human lineage. Distance to the nearest possible Homo erectus remains for Southeast and East Asia and the westward located Levantine, African and European remains are no less than exorbitant three to four thousand kilometres from the Hathnora fossil locality. The nearest possible 25 000–30 000-year-old descendents of Batadombalena (Sri Lanka), Darra-i-Kur (Afghanistan), Kurnool District caves, or even younger Bhimbetka, Sarai Nahar, Mahadaha, Lothal, Dholavira of 4000–10 000-year-old antiquity are closely located (Fig. 1). A possible hierarchical relationship is attempted. To cite this article: A. Sonakia, H. de Lumley, C. R. Palevol 5 (2006).
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Background: Recent analyses of de novo DNA mutations in modern humans have suggested a nuclear substitution rate that is approximately half that of previous estimates based on fossil calibration. This result has led to suggestions that major events in human evolution occurred far earlier than previously thought. Results: Here, we use mitochondrial genome sequences from ten securely dated ancient modern humans spanning 40,000 years as calibration points for the mitochondrial clock, thus yielding a direct estimate of the mitochondrial substitution rate. Our clock yields mitochondrial divergence times that are in agreement with earlier estimates based on calibration points derived from either fossils or archaeological material. In particular, our results imply a separation of non-Africans from the most closely related sub-Saharan African mitochondrial DNAs (haplogroup L3) that occurred less than 62-95 kya. Conclusions: Though single loci like mitochondrial DNA (mtDNA) can only provide biased estimates of population divergence times, they can provide valid upper bounds. Our results exclude most of the older dates for African and non-African population divergences recently suggested by de novo mutation rate estimates in the nuclear genome.
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To eliminate possible confusion in the reporting of isotopic abundances on non-corresponding scales, the Commission on Atomic Weights and Isotopic Abundances recommended at the 37 th General Assembly at Lisbon, Portugal that (i) 2 H/ 1 H relative ratios of all substances be expressed relative to VSMOW (Vienna Standard Mean Ocean Water) on a scale such that 2 H/ 1 H of SLAP (Standard Light Antarctic Precipitation) is 0.572 times that of VSMOW, (ii) 13 C/ 12 C relative ratios of all substances be expressed relative to VPDB (Vienna Peedee belemnite) on a scale such that 13 C/ 12 C of NBS 19 carbonate is 1.00195 times that of VPDB, and (iii) 18 O/ 16 O ratios of all substances be expressed relative to either VSMOW or VPDB on scales such that 18 O/ 16 O of SLAP is 0.9445 times that of VSMOW.
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One hundred and forty middle ear ossicles belonging to three different populations were examined. General homogeneity was observed in these groups of modern man, in spite of their different ethnic and geographic origins as well as chronological age.
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Perceptions of the emergence and spread of modern humans have changed recently through the reanalysis of fossils, an improved geochronological framework, and the discovery of a few specimens. Early modern humans in various portions of the Old World exhibit complex and varying mosaics of archaic, modern, and regional mor-phological characteristics. On the basis of this pattern, in conjunc-tion with the emerging chronology of the earliest modern humans, the paleontological data indicate an assimilation model for modern human origins, in which the earliest modern humans emerged in eastern Africa, dispersed briefly into southwestern Asia, and then subsequently spread into the remainder of Africa and southern Asia, eventually into higher latitude Eurasia. The earliest modern humans outside of the core area of eastern Africa can be understood only if a variable degree of admixture with regional groups of late archaic hu-mans occurred. Current and expected fossil and molecular data are unlikely to illuminate the degree of assimilation that took place in most regions of the Old World. However, the current chronological and phylogenetic framework provides the basis for ongoing investi-gation of the nature of this Late Pleistocene transitional period.
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The recovery of DNA sequences from early modern humans (EMHs) could shed light on their interactions with archaic groups such as Neandertals and their relationships to current human populations. However, such experiments are highly problematic because present-day human DNA frequently contaminates bones [1, 2]. For example, in a recent study of mitochondrial (mt) DNA from Neolithic European skeletons, sequence variants were only taken as authentic if they were absent or rare in the present population, whereas others had to be discounted as possible contamination [3, 4]. This limits analysis to EMH individuals carrying rare sequences and thus yields a biased view of the ancient gene pool. Other approaches of identifying contaminating DNA, such as genotyping all individuals who have come into contact with a sample, restrict analyses to specimens where this is possible [5, 6] and do not exclude all possible sources of contamination. By studying mtDNA in Neandertal remains, where contamination and endogenous DNA can be distinguished by sequence, we show that fragmentation patterns and nucleotide misincorporations can be used to gauge authenticity of ancient DNA sequences. We use these features to determine a complete mtDNA sequence from a approximately 30,000-year-old EMH from the Kostenki 14 site in Russia.
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A complete mitochondrial (mt) genome sequence was reconstructed from a 38,000 year-old Neandertal individual with 8341 mtDNA sequences identified among 4.8 Gb of DNA generated from approximately 0.3 g of bone. Analysis of the assembled sequence unequivocally establishes that the Neandertal mtDNA falls outside the variation of extant human mtDNAs, and allows an estimate of the divergence date between the two mtDNA lineages of 660,000 +/- 140,000 years. Of the 13 proteins encoded in the mtDNA, subunit 2 of cytochrome c oxidase of the mitochondrial electron transport chain has experienced the largest number of amino acid substitutions in human ancestors since the separation from Neandertals. There is evidence that purifying selection in the Neandertal mtDNA was reduced compared with other primate lineages, suggesting that the effective population size of Neandertals was small.
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In 1997 a human mandibular second deciduous incisor was discovered during excavations at the central Asian Middle Paleolithic site of Khudji, Tajikistan. The specimen was associated with a late Middle Paleolithic assemblage in a minimally disturbed cultural layer. The specimen is average in size for Late Pleistocene archaic human di(2)s and differs from many late archaic human di(2)s in having minimal marginal ridges and tapering markedly distally. In these features it resembles a minority of specimens from the later Pleistocene.
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The temporal bone is the location of several traits thought to differentiate Neanderthals from modern humans, including some proposed Neanderthal-derived traits. Most of these, however, are difficult to measure and are usually described qualitatively. This study applied the techniques of geometric morphometrics to the complex morphology of the temporal bone, in order to quantify the differences observed between Neanderthal and modern human anatomy. Two hundred and seventy modern human crania were measured, representing 9 populations of 30 individuals each, and spanning the extremes of the modern human geographical range. Twelve Neanderthal specimens, as well as Reilingen, Kabwe, Skhul 5, Qafzeh 9, and 4 Late Paleolithic European specimens, were included in the fossil sample. The data were collected in the form of three-dimensional (3-D) landmark coordinates, and specimen configurations were superimposed using generalized Procrustes analysis. The fitted coordinates were then analyzed by an array of multivariate statistical methods, including principal components analysis, canonical variates analysis, and Mahalanobis D(2). The temporal bone landmark analysis was very successful in separating Neanderthals from modern humans. Neanderthals were separated from modern humans in both the principal components and canonical variates analyses. They were much further in Mahalanobis distances from all modern human populations than any two modern human groups were from each other. Most of the previously described temporal bone traits contributed to this separation.
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This paper presents a comprehensive comparative analysis of the Neanderthal bony labyrinth, a structure located inside the petrous temporal bone. Fifteen Neanderthal specimens are compared with a Holocene human sample, as well as with a small number of European Middle Pleistocene hominins, and early anatomically modern and European Upper Palaeolithic humans. Compared with Holocene humans the bony labyrinth of Neanderthals can be characterized by an anterior semicircular canal arc which is smaller in absolute and relative size, is relatively narrow, and shows more torsion. The posterior semicircular canal arc is smaller in absolute and relative size as well, it is more circular in shape, and is positioned more inferiorly relative to the lateral canal plane. The lateral semicircular canal arc is absolutely and relatively larger. Finally, the Neanderthal ampullar line is more vertically inclined relative to the planar orientation of the lateral canal. The European Upper Palaeolithic and early modern humans are most similar, although not fully identical to Holocene humans in labyrinthine morphology. The European Middle Pleistocene hominins show the typical semicircular canal morphology of Neanderthals, with the exception of the arc shape and inferiorly position of the posterior canal and the strongly inclined ampullar line. The marked difference between the labyrinths of Neanderthals and modern humans can be used to assess the phylogenetic affinities of fragmentary temporal bone fossils. However, this application is limited by a degree of overlap between the morphologies. The typical shape of the Neanderthal labyrinth appears to mirror aspects of the surrounding petrous pyramid, and both may follow from the phylogenetic impact of Neanderthal brain morphology moulding the shape of the posterior cranial fossa. The functionally important arc sizes of the Neanderthal semicircular canals may reflect a pattern of head movements different from that of modern humans, possibly related to aspects of locomotor behaviour and the kinematic properties of their head and neck.