Результаты исследования древней ДНК (дДНК)

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Genomic Evidence Establishes Anatolia as the Source of the European Neolithic Gene Pool

Ayça Omrak, Torsten Günther, Cristina Valdiosera, Emma M. Svensson, Helena Malmström, Henrike Kiesewetter, William Aylward, Jan Storå, Mattias Jakobsson, Anders Götherström

Accepted: December 9, 2015
Received in revised form: December 7, 2015
Received: November 20, 2015

Summary

Anatolia and the Near East have long been recognized as the epicenter of the Neolithic expansion through archaeological evidence. Recent archaeogenetic studies on Neolithic European human remains have shown that the Neolithic expansion in Europe was driven westward and northward by migration from a supposed Near Eastern origin [ 1–5 ]. However, this expansion and the establishment of numerous culture complexes in the Aegean and Balkans did not occur until 8,500 before present (BP), over 2,000 years after the initial settlements in the Neolithic core area [ 6–9 ]. We present ancient genome-wide sequence data from 6,700-year-old human remains excavated from a Neolithic context in Kumtepe, located in northwestern Anatolia near the well-known (and younger) site Troy [ 10 ]. Kumtepe is one of the settlements that emerged around 7,000 BP, after the initial expansion wave brought Neolithic practices to Europe. We show that this individual displays genetic similarities to the early European Neolithic gene pool and modern-day Sardinians, as well as a genetic affinity to modern-day populations from the Near East and the Caucasus. Furthermore, modern-day Anatolians carry signatures of several admixture events from different populations that have diluted this early Neolithic farmer component, explaining why modern-day Sardinian populations, instead of modern-day Anatolian populations, are genetically more similar to the people that drove the Neolithic expansion into Europe. Anatolia’s central geographic location appears to have served as a connecting point, allowing a complex contact network with other areas of the Near East and Europe throughout, and after, the Neolithic.

http://www.cell.com/current-biology/abstract/S0960-9822(15)01516-X

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Iron Age and Anglo-Saxon genomes from East England reveal British migration history

Stephan Schiffels, Wolfgang Haak, Pirita Paajanen, Bastien Llamas, Elizabeth Popescu, Louise Loe, Rachel Clarke, Alice Lyons, Richard Mortimer, Duncan Sayer, Chris Tyler-Smith, Alan Cooper & Richard Durbin.

Abstract

British population history has been shaped by a series of immigrations, including the early Anglo-Saxon migrations after 400 CE. It remains an open question how these events affected the genetic composition of the current British population. Here, we present whole-genome sequences from 10 individuals excavated close to Cambridge in the East of England, ranging from the late Iron Age to the middle Anglo-Saxon period. By analysing shared rare variants with hundreds of modern samples from Britain and Europe, we estimate that on average the contemporary East English population derives 38% of its ancestry from Anglo-Saxon migrations. We gain further insight with a new method, rarecoal, which infers population history and identifies fine-scale genetic ancestry from rare variants. Using rarecoal we find that the Anglo-Saxon samples are closely related to modern Dutch and Danish populations, while the Iron Age samples share ancestors with multiple Northern European populations including Britain.

http://www.nature.com/ncomms/2016/160119/ncomms10408/full/ncomms10408.html

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Genomic signals of migration and continuity in Britain before the Anglo-Saxons

Rui Martiniano, Anwen Caffell, Malin Holst, Kurt Hunter-Mann, Janet Montgomery,Gundula Müldner, Russell L. McLaughlin, Matthew D. Teasdale, Wouter van Rheenen, Jan H. Veldink, Leonard H. van den Berg, Orla Hardiman, Maureen Carroll,    Steve Roskams, John Oxley, Colleen Morgan, Mark G. Thomas, Ian Barnes, Christine McDonnell,   Matthew J. Collins et al.

Abstract

The purported migrations that have formed the peoples of Britain have been the focus of generations of scholarly controversy. However, this has not benefited from direct analyses of ancient genomes. Here we report nine ancient genomes (~1 ×) of individuals from northern Britain: seven from a Roman era York cemetery, bookended by earlier Iron-Age and later Anglo-Saxon burials. Six of the Roman genomes show affinity with modern British Celtic populations, particularly Welsh, but significantly diverge from populations from Yorkshire and other eastern English samples. They also show similarity with the earlier Iron-Age genome, suggesting population continuity, but differ from the later Anglo-Saxon genome. This pattern concords with profound impact of migrations in the Anglo-Saxon period. Strikingly, one Roman skeleton shows a clear signal of exogenous origin, with affinities pointing towards the Middle East, confirming the cosmopolitan character of the Empire, even at its northernmost fringes.

http://www.nature.com/ncomms/2016/160119/ncomms10326/full/ncomms10326.html

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A Community in Life and Death: The Late Neolithic Megalithic Tomb at Alto de Reinoso (Burgos, Spain)

Kurt W. Alt  , Stephanie Zesch , Rafael Garrido-Pena, Corina Knipper, Anna Szécsényi-Nagy, Christina Roth, Cristina Tejedor-Rodríguez, Petra Held, Íñigo García-Martínez-de-Lagrán, Denise Navitainuck, Héctor Arcusa Magallón, Manuel A. Rojo-Guerra

Published: January 20, 2016

Abstract

The analysis of the human remains from the megalithic tomb at Alto de Reinoso represents the widest integrative study of a Neolithic collective burial in Spain. Combining archaeology, osteology, molecular genetics and stable isotope analysis (87Sr/86Sr, δ15N, δ13C) it provides a wealth of information on the minimum number of individuals, age, sex, body height, pathologies, mitochondrial DNA profiles, kinship relations, mobility, and diet. The grave was in use for approximately one hundred years around 3700 cal BC, thus dating from the Late Neolithic of the Iberian chronology. At the bottom of the collective tomb, six complete and six partial skeletons lay in anatomically correct positions. Above them, further bodies represented a subsequent and different use of the tomb, with almost all of the skeletons exhibiting signs of manipulation such as missing skeletal parts, especially skulls. The megalithic monument comprised at least 47 individuals, including males, females, and subadults, although children aged 0–6 years were underrepresented. The skeletal remains exhibited a moderate number of pathologies, such as degenerative joint diseases, healed fractures, cranial trauma, and a low intensity of caries. The mitochondrial DNA profiles revealed a pattern pointing to a closely related local community with matrilineal kinship patterns. In some cases adjacent individuals in the bottom layer showed familial relationships. According to their strontium isotope ratios, only a few individuals were likely to have spent their early childhood in a different geological environment, whilst the majority of individuals grew up locally. Carbon and nitrogen isotope analysis, which was undertaken to reconstruct the dietary habits, indicated that this was a homogeneous group with egalitarian access to food. Cereals and small ruminants were the principal sources of nutrition. These data fit in well with a lifestyle typical of sedentary farming populations in the Spanish Meseta during this period of the Neolithic.

http://journals.plos.org/plosone/article?id=10.1371/journal.pone.0146176

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United in death—related by blood? Genetic and archeometric analyses of skeletal remains from the neolithic earthwork bruchsal-aue

Marcel Keller, Andreas Rott, Nadja Hoke, Heiner Schwarzberg, Birgit Regner-Kamlah, Michaela Harbeck andJoachim Wahl

Article first published online: 25 MAR 2015

ABSTRACT

Objectives: Straight next to a segment of the outer ditch of the Late Neolithic Michelsberg Culture earthwork of Bruchsal-Aue in SW-Germany (ca. 4250–3650 calBC), a multiple burial of eight individuals (two male adults and six children) plus a subsequent child burial was excavated. In this study, we applied a multidisciplinary approach to elucidate interpersonal relationships and life histories within this collective. Materials and methods: To determine the identity of this collective, we performed aDNA analyses in addition to osteological examination using HVR I plus Y-chromosomal and autosomal STR profiling to find evidence for kinship relations. Strontium isotopic analyses were used to reconsider migrational behavior. To find evidence for a specific social affiliation, the individual diet was reconstructed by performing nitrogen and carbon isotopic analyses. Furthermore, radiocarbon-dating was carried out to integrate the burial context into an absolute timeframe. Two nearby single burials were included in the analyses for comparison. Results: Because of a shared HVR I haplotype, three pairs of individuals were most likely linked by kinship, and statistical testing on autosomal STR profiles shows a high probability for the pair of two men being brothers. Although it cannot be excluded, isotopic data gave no clear proof for migration. A rather poor health status is indicated by skeletal stress markers even though the isotope data attest to a diet rich in meat and fish. Discussion: Although clear kinship relations among the infants remain unconfirmed, a relationship could also be indicated by the positioning of the bodies in the burial pit. Whereas a common cause of death might have been the presupposition for their special treatment, interpersonal relationships were likely the decisive factor for the multiple burial.

http://onlinelibrary.wiley.com/doi/10.1002/ajpa.22738/abstract

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Pleistocene Mitochondrial Genomes Suggest a Single Major Dispersal of Non-Africans and a Late Glacial Population Turnover in Europe

Cosimo Posth, Gabriel Renaud, Alissa Mittnik, Dorothée G. Drucker, Hélène Rougier, Christophe Cupillard, Frédérique Valentin, Corinne Thevenet, Anja Furtwängler, Christoph Wißing, Michael Francken, Maria Malina, Michael Bolus and all.

Summary

How modern humans dispersed into Eurasia and Australasia, including the number of separate expansions and their timings, is highly debated [ 1, 2 ]. Two categories of models are proposed for the dispersal of non-Africans: (1) single dispersal, i.e., a single major diffusion of modern humans across Eurasia and Australasia [ 3–5 ]; and (2) multiple dispersal, i.e., additional earlier population expansions that may have contributed to the genetic diversity of some present-day humans outside of Africa [ 6–9 ]. Many variants of these models focus largely on Asia and Australasia, neglecting human dispersal into Europe, thus explaining only a subset of the entire colonization process outside of Africa [ 3–5, 8, 9 ]. The genetic diversity of the first modern humans who spread into Europe during the Late Pleistocene and the impact of subsequent climatic events on their demography are largely unknown. Here we analyze 55 complete human mitochondrial genomes (mtDNAs) of hunter-gatherers spanning ∼35,000 years of European prehistory. We unexpectedly find mtDNA lineage M in individuals prior to the Last Glacial Maximum (LGM). This lineage is absent in contemporary Europeans, although it is found at high frequency in modern Asians, Australasians, and Native Americans. Dating the most recent common ancestor of each of the modern non-African mtDNA clades reveals their single, late, and rapid dispersal less than 55,000 years ago. Demographic modeling not only indicates an LGM genetic bottleneck, but also provides surprising evidence of a major population turnover in Europe around 14,500 years ago during the Late Glacial, a period of climatic instability at the end of the Pleistocene.

http://www.cell.com/current-biology/fulltext/S0960-9822%2816%2900087-7

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Identifications of Ancient Egyp tian Royal Mummies from the 18th Dynasty Reconsidered

M.E. Habicht, A.S. Bouwman, and F.J.

Institute for Evolutionary Medicine, University of Zurich, CH 8057, Switzerland

ABSTRACT
For centuries, ancient Egyptian Royal mummies have drawn the attention both of the general public and scien-
tists. Many royal mummies from the New Kingdom have survived. The discoveries of the bodies of these ancient rul-
ers have always sparked much attention, yet not all identifications are clear even nowadays. This study presents a
meta-analysis to demonstrate the difficulties in identifying ancient Egyptian royal mummies. Various methods and
pitfalls in the identification of the Pharaohs are reassessed since new scientific methods can be used, such as ancient
DNA-profiling and CT-scanning. While the ancestors of Tutankhamun have been identified, some identities are still
highly controversial (e.g., the mystery of the KV-55 skeleton, recently most likely identified as the genetic father of
Tutankhamun). The meta-analysis confirms the suggested identity of some mummies (e.g., Amenhotep III, Thutmo-
sis IV, and Queen Tjye). Am J Phys Anthropol 159:S216–S231, 2016.

http://onlinelibrary.wiley.com/doi/10.1002/ajpa.22909/epdf

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Standing upright to all eternity – The Mesolithic burial site at Groß Fredenwalde, Brandenburg (NE Germany)

Aufrecht in die Ewigkeit – Der mesolithische Bestatt

Thomas Terberger, Andreas Kotula, Sebastian Lorenz, Manuela Schult, Joachim Burger.

Abstract

This article presents results of new research on the Mesolithic burial site at Groß Fredenwalde in northeastern Germany, where a multiple burial was first discovered by accident in 1962. Anthropological analyses identified one female with a child and two males with two children within this material. According to systematic AMS dating and
C-isotope analyses the individuals are typical Mesolithic hunter-fisher-gatherers of the Atlantic period (c. 6
000 calBC). During re-excavation of the site in 2012-2014 three new burials including a disturbed child burial and a baby burial were recognised. There is also an outstanding burial: a young man was interred standing upright and then furnished in stages. The burial is without any parallel in Central Europe, although there are possible parallels at Olenij Ostrov in Karelia. Altogether nine individuals from at least four graves are now known; they probably belong to an early cemetery located at a prominent position in the landscape. AMS-dates assign the burials to the period from c. 6400 to 4900 calBC, and thus the site was in use when the first Linear Band Pottery farmers established the gricultural way of life in the region c. 5200 calBC. Two successfully analysed individuals belong to the haplogroup U of mitochondrial lineages fitting well into the model of highly differentiated forager and farmer populations.

http://www.quartaer.eu/pdfs/2015/2015_06_terberger.pdf

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Early Medieval Muslim Graves in France: First Archaeological, Anthropological and Palaeogenomic Evidence

Yves Gleize, Fanny Mendisco, Marie-Hélène Pemonge, Christophe Hubert, Alexis Groppi, Bertrand Houix, Marie-France Deguilloux, Jean-Yves Breuil.

Abstract

The rapid Arab-Islamic conquest during the early Middle Ages led to major political and cultural changes in the Mediterranean world. Although the early medieval Muslim presence in the Iberian Peninsula is now well documented, based in the evaluation of archeological and historical sources, the Muslim expansion in the area north of the Pyrenees has only been documented so far through textual sources or rare archaeological data. Our study provides the first archaeo-anthropological testimony of the Muslim establishment in South of France through the multidisciplinary analysis of three graves excavated at Nimes. First, we argue in favor of burials that followed Islamic rites and then note the presence of a community practicing Muslim traditions in Nimes. Second, the radiometric dates obtained from all three human skeletons (between the 7th and the 9th centuries AD) echo historical sources documenting an early Muslim presence in southern Gaul (i.e., the first half of 8th century AD). Finally, palaeogenomic analyses conducted on the human remains provide arguments in favor of a North African ancestry of the three individuals, at least considering the paternal lineages. Given all of these data, we propose that the skeletons from the Nimes burials belonged to Berbers integrated into the Umayyad army during the Arab expansion in North Africa. Our discovery not only discusses the first anthropological and genetic data concerning the Muslim occupation of the Visigothic territory of Septimania but also highlights the complexity of the relationship between the two communities during this period.
http://journals.plos.org/plosone/article?id=10.1371/journal.pone.0148583

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В.В. Питулько, В. И. Хартанович, В. Б. Тимошин, В. Г. Часнык, Е. Ю. Павлова, А. К. Каспаров
ДРЕВНЕЙШИЕ АНТРОПОЛОГИЧЕСКИЕ НАХОДКИ ВЫСОКОШИРОТНОЙ АРКТИКИ
(Жоховская стоянка, Новосибирские острова)*

В ходе раскопок Жоховской стоянки (2000–2005 гг.), расположенной под 76° с. ш. в Сибирской Арктике на острове Жохова в архипелаге Де Лонга, получены антропологические материалы, являющиеся древнейшими находками подобного рода в Арктике (~8000 л. н.). Исследование выделенной мтДНК позволило установить пол и степень родства изученных образцов. В большинстве случаев они принадлежат к гаплогруппе К. Данная группа, имеющая западноевразийские корни, предшествовала в арктической Сибири современным этносам (включая юкагиров) и была замещена ими во второй половине голоцена.

http://uralhist.uran.ru/pdf/Pitulko_all.pdf

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Ancient mitochondrial DNA provides high-resolution time scale of the peopling of the Americas

Bastien Llamas1,*,†, Lars Fehren-Schmitz2,†, Guido Valverde1, Julien Soubrier1, Swapan Mallick3,4,5, Nadin Rohland3,4,5, Susanne Nordenfelt3,4,5, Cristina Valdiosera6, Stephen M. Richards1, Adam Rohrlach7, Maria Inés Barreto Romero8, Isabel Flores Espinoza8, Elsa Tomasto Cagigao9, Lucía Watson Jiménez9,10, Krzysztof Makowski9, Ilán Santiago Leboreiro Reyna11, Josefina Mansilla Lory11, Julio Alejandro Ballivián Torrez12, Mario A. Rivera13, Richard L. Burger14, Maria Constanza Ceruti15,16, Johan Reinhard17, R. Spencer Wells17,‡, Gustavo Politis18, Calogero M. Santoro19, Vivien G. Standen19, Colin Smith6, David Reich3,4,5, Simon Y. W. Ho20, Alan Cooper1,*,§ and Wolfgang Haak1,*

Abstract

The exact timing, route, and process of the initial peopling of the Americas remains uncertain despite much research. Archaeological evidence indicates the presence of humans as far as southern Chile by 14.6 thousand years ago (ka), shortly after the Pleistocene ice sheets blocking access from eastern Beringia began to retreat. Genetic estimates of the timing and route of entry have been constrained by the lack of suitable calibration points and low genetic diversity of Native Americans. We sequenced 92 whole mitochondrial genomes from pre-Columbian South American skeletons dating from 8.6 to 0.5 ka, allowing a detailed, temporally calibrated reconstruction of the peopling of the Americas in a Bayesian coalescent analysis. The data suggest that a small population entered the Americas via a coastal route around 16.0 ka, following previous isolation in eastern Beringia for ~2.4 to 9 thousand years after separation from eastern Siberian populations. Following a rapid movement throughout the Americas, limited gene flow in South America resulted in a marked phylogeographic structure of populations, which persisted through time. All of the ancient mitochondrial lineages detected in this study were absent from modern data sets, suggesting a high extinction rate. To investigate this further, we applied a novel principal components multiple logistic regression test to Bayesian serial coalescent simulations. The analysis supported a scenario in which European colonization caused a substantial loss of pre-Columbian lineages.

http://advances.sciencemag.org/content/2/4/e1501385.full

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The genetic history of Ice Age Europe

Qiaomei Fu,   Cosimo Posth,   Mateja Hajdinjak,   Martin Petr,   Swapan Mallick,   Daniel Fernandes,   Anja Furtwängler,   Wolfgang Haak,   Matthias Meyer,   Alissa Mittnik,   Birgit Nickel,   Alexander Peltzer,   Nadin Rohland,   Viviane Slon,   Sahra Talamo,   Iosif Lazaridis,   Mark Lipson,   Iain Mathieson,   Stephan Schiffels,   Pontus Skoglund,   Anatoly P. Derevianko,   Nikolai Drozdov,   Vyacheslav Slavinsky,   Alexander Tsybankov,   Renata Grifoni Cremonesi   et al.

Published online 02 May 2016

Abstract

Modern humans arrived in Europe ~45,000 years ago, but little is known about their genetic composition before the start of farming ~8,500 years ago. Here we analyse genome-wide data from 51 Eurasians from ~45,000–7,000 years ago. Over this time, the proportion of Neanderthal DNA decreased from 3–6% to around 2%, consistent with natural selection against Neanderthal variants in modern humans. Whereas there is no evidence of the earliest modern humans in Europe contributing to the genetic composition of present-day Europeans, all individuals between ~37,000 and ~14,000 years ago descended from a single founder population which forms part of the ancestry of present-day Europeans. An ~35,000-year-old individual from northwest Europe represents an early branch of this founder population which was then displaced across a broad region, before reappearing in southwest Europe at the height of the last Ice Age ~19,000 years ago. During the major warming period after ~14,000 years ago, a genetic component related to present-day Near Easterners became widespread in Europe. These results document how population turnover and migration have been recurring themes of European prehistory.

http://www.nature.com/nature/journal/vaop/ncurrent/full/nature17993.html#access

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Mitochondrial DNA Reveals the Trace of the Ancient Settlers of a Violently Devastated Late Bronze and Iron Ages Village

Carolina Núñez, Miriam Baeta, Sergio Cardoso, Leire Palencia-Madrid, Noemí García-Romero, Armando Llanos,   Marian M. de Pancorbo

PLOS Published: May 13, 2016 http://dx.doi.org/10.1371/journal.pone.0155342

Abstract

La Hoya (Alava, Basque Country) was one of the most important villages of the Late Bronze and Iron Ages of the north of the Iberian Peninsula, until it was violently devastated around the 4th century and abandoned in the 3rd century B.C. Archaeological evidences suggest that descendants from La Hoya placed their new settlement in a nearby hill, which gave rise to the current village of Laguardia. In this study, we have traced the genetic imprints of the extinct inhabitants of La Hoya through the analysis of maternal lineages. In particular, we have analyzed the mitochondrial DNA (mtDNA) control region of 41 human remains recovered from the archaeological site for comparison with a sample of 51 individuals from the geographically close present-day population of Laguardia, as well as 56 individuals of the general population of the province of Alava, where the archaeological site and Laguardia village are located. MtDNA haplotypes were successfully obtained in 25 out of 41 ancient samples, and 14 different haplotypes were identified. The major mtDNA subhaplogroups observed in La Hoya were H1, H3, J1 and U5, which show a distinctive frequency pattern in the autochthonous populations of the north of the Iberian Peninsula. Approximate Bayesian Computation analysis was performed to test the most likely model for the local demographic history. The results did not sustain a genealogical continuity between Laguardia and La Hoya at the haplotype level, although factors such as sampling effects, recent admixture events, and genetic bottlenecks need to be considered. Likewise, the highly similar subhaplogroup composition detected between La Hoya and Laguardia and Alava populations do not allow us to reject a maternal genetic continuity in the human groups of the area since at least the Iron Age to present times. Broader analyses, based on a larger collection of samples and genetic markers, would be required to study fine-scale population events in these human groups.

http://journals.plos.org/plosone/article?id=10.1371%2Fjournal.pone.0155342

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A European Mitochondrial Haplotype Identified in Ancient Phoenician Remains from Carthage, North Africa

Elizabeth A. Matisoo-Smith, Anna L. Gosling, James Boocock, Olga Kardailsky, Yara Kurumilian, Sihem Roudesli-Chebbi, Leila Badre, Jean-Paul Morel, Leïla Ladjimi Sebaï, Pierre A. Zalloua

PLOS Published: May 25, 2016

Abstract

While Phoenician culture and trade networks had a significant impact on Western civilizations, we know little about the Phoenicians themselves. In 1994, a Punic burial crypt was discovered on Byrsa Hill, near the entry to the National Museum of Carthage in Tunisia. Inside this crypt were the remains of a young man along with a range of burial goods, all dating to the late 6th century BCE. Here we describe the complete mitochondrial genome recovered from the Young Man of Byrsa and identify that he carried a rare European haplogroup, likely linking his maternal ancestry to Phoenician influenced locations somewhere on the North Mediterranean coast, the islands of the Mediterranean or the Iberian Peninsula. This result not only provides the first direct ancient DNA evidence of a Phoenician individual but the earliest evidence of a European mitochondrial haplogroup, U5b2c1, in North Africa.

http://journals.plos.org/plosone/article?id=10.1371/journal.pone.0155046

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The mitogenome of a 35,000-year-old Homo sapiens from Europe supports a Palaeolithic back-migration to Africa

M. Hervella, E. M. Svensson, A. Alberdi, T. Günther, N. Izagirre, A. R. Munters, S. Alonso, M. Ioana, F. Ridiche, A. Soficaru, M. Jakobsson, M. G. Netea & C. de-la-Rua

Scientific Reports 6, Article number: 25501 (2016) doi:10.1038/srep25501
   
Received: 15 February 2016 Аccepted: 19 April 2016 Published online: 19 May 2016

Abstract

After the dispersal of modern humans (Homo sapiens) Out of Africa, hominins with a similar morphology to that of present-day humans initiated the gradual demographic expansion into Eurasia. The mitogenome (33-fold coverage) of the Peştera Muierii 1 individual (PM1) from Romania (35 ky cal BP) we present in this article corresponds fully to Homo sapiens, whilst exhibiting a mosaic of morphological features related to both modern humans and Neandertals. We have identified the PM1 mitogenome as a basal haplogroup U6*, not previously found in any ancient or present-day humans. The derived U6 haplotypes are predominantly found in present-day North-Western African populations. Concomitantly, those found in Europe have been attributed to recent gene-flow from North Africa. The presence of the basal haplogroup U6* in South East Europe (Romania) at 35 ky BP confirms a Eurasian origin of the U6 mitochondrial lineage. Consequently, we propose that the PM1 lineage is an offshoot to South East Europe that can be traced to the Early Upper Paleolithic back migration from Western Asia to North Africa, during which the U6 lineage diversified, until the emergence of the present-day U6 African lineages.

http://journals.plos.org/plosone/article?id=10.1371/journal.pone.0155046