RYM KÉFI, ALAIN STEVANOVITCH, ERIC BOUZAID, ELIANE BÉRAUD-COLOMB
DIVERSITÉ MITOCHONDRIALE DE LA POPULATION DE TAFORALT (12.000 ANS BP – MAROC): UNE APPROCHE GÉNÉTIQUE À L'ÉTUDE DU PEUPLEMENT DE L'AFRIQUE DU NORD 2005
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excerpts, translated to English
MITOCHONDRIAL DIVERSITY OF THE POPULATION Taforalt (12,000 YEARS BP - MOROCCO): GENETIC APPROACH TO THE STUDY OF SETTLEMENT NORTH AFRICA
MITOCHONDRIAL DIVERSITY OF THE POPULATION Taforalt (12,000 YEARS BP - MOROCCO): GENETIC APPROACH TO THE STUDY OF SETTLEMENT NORTH AFRICA SUMMARY: anthropological and genetic studies have revealed the complexity of the settlement of North Africa. In order to contribute to a better understanding of the development of people in the Mediterranean region, we proposed to study mitochondrial diversity of the population ibéromaurusienne Taforalt (12,000 years - Morocco). The ancient DNA was extracted thirty specimens unearthed the cave épipaléolithique Taforalt. We amplified and sequenced the HVS1 fragment of the mitochondrial DNA control region. The genetic diversity of the population Taforalt shows that it is not related individuals to sub-Saharan individuals. The hypothesis of a sub-Sudanese ibéromaurusiens of men is rejected. Our results are in line with local origins of the population Taforalt and genetic continuity in North Africa. KEYWORDS: ancient DNA - mitochondrial DNA - iberomaurusian - Epipalaeolithic - Taforalt - North Africa Mitochondrial DNA DIVERSITY OF POPULATION Taforalt (12,000 YEARS BP, MOROCCO): A GENETIC STUDY OF THE SETTLEMENT OF NORTH AFRICA ABSTRACT: The people exhumed from the archaeological site of Taforalt in Morocco (12,000 years BP) is a valuable source of information Toward a better knowledge of the settlement of Northern Africa Region and Provides a revolutionary way to Specify the origin of Ibero-Maurusian populations. Was Ancient DNA Extracted from 31 bone remains from Taforalt. The HVS1 fragment of the mitochondrial DNA control PCR-amplified Region Was Directly and sequenced. Mitochondrial diversity in Taforalt shows the lack of sub-Saharan haplogroups Suggesting That Ibero-Maurusian Individuals HAD not originated in sub-Saharan area. Our results reveal a probable local Evolution of Taforalt population and a genetic continuity in North Africa. KEY WORDS: Ancient DNA - mitochondrial DNA polymorphism - Ibero-Maurusian - Epipaleolithic - Taforalt - North Africa. INTRODUCTION North Africa stand of knowledge is construitsuivantdeuxapproches: the uneàtraversl'étudedes archaeological specimens and their environment, the other through the study of the molecular diversity of different genetic markers worn by individuals in the current population. The markers are studied 1 Rym Kefi Stevanovitch Alain, Eric Bouzaid, Eliane Beraud-Colomb a) b) FIGURE 1. 1a: Overview of the cave Taforalt. 1b specimen Skull Taf VIII. Mitochondrial DNA, the Y chromosome, the HLA system, hemoglobin and certain microsatellites, to name the main ones. The oldest human presence in North Africa date of 700,000 years BP and corresponds to the skeletal remains of mauretanicus Atlanthropus. This Homo erectus was discovered in Ternifine in Algeria (Arambourg 1963). Specimens of Sidi Abdderrahman sites (200,000 years BP), Jebel Irhoud (100,000 years BP), Ain Hanech, Salé, Rabat (160,000 BP) and Dar es Soltan (70,000 years BP, morphologically similar to modern Homo sapiens sapiens ) Morocco (Camps 1974) could witness the transition from Homo erectus to Homo sapiens sapiens. The lithic industry associated with north African Homo erectus was the Middle Paleolithic (Mousterian and Levallois industries). With the emergence of Homo sapiens, an original industry had developed: Aterian specific western North Africa (Morocco, Algeria and Tunisia) (Camps 1974 DEBENATH et al. 1986). Highlighted on the sites of Dar es Soltan and Taforalt (skull fragments in a lower layer of the site Taforalt) (Ferembach 1962), Human Aterian this morphologically modern characters but also preserves more archaic characteristics than those men Crô-Magnon. It also has enough homology with human moustérien Jebel Irhoud, for us to consider a relationship between them (Camps 1989). Finally this man Aterian could be the ancestor of Human Mechta El-Arbi (geographical successor to 20,000 years BP). However the question remains open. Various origins are cited to explain the onset of Human Mechta el-Arbi, author of ibéromaurusienne industry in the Maghreb. It could be coming to Europe via Spain, although no milestone has been found between southern France and the Strait of Gibraltar (Camps 1989). It could also be from Palestine (Middle East) and come from a common center from which would be developed two branches: one to Europe giving human Crô-Magnon, and the other to the Maghreb, giving human Mechta El Arbi. The latter theory, based on morphological comparisons between populations cromagnoïdes and men of Mechta el-Arbi, is defended by several authors (Vallois, Movius 1952 Balout 1954). The presence of Proto Crô-Magnons (Homo sapiens sapiens) in the Middle East (and Qafzeh Skhul sites, dated to 100,000 and 92,000 years BP) reinforces this hypothesis. Other authors suggest an origin of man of Mechta El-Arbi in Nubia. Indeed, many similarities were found between the Nubians of the late Pleistocene and men of Mechta El-Arbi. The Nubian Microlithic industry Qadan (15,000 to 11,000 years BP) is similar to the industry ibéromaurusienne Maghreb. neighboring burial practices can be observed, including the site of Jebel Sahaba in Lower Nubia (Wendorf 1968). Moreover, many ibéromaurusiens skeletons, such as Dar es Soltan and Taforalt Morocco or Ali Bacha Afalou and Algeria, have similarities with people of 12,000 years of Wadi Halfa and Jebal Sahaba Nubia (Wendorf 1968) . Similarities are also observed with individuals of Nazlet Khater and those of Wadi Kubbaniya in Egypt, dating back to 25,000 years BP (Wendorf 1968). Regarding the genetic approach of the settlement of North Africa, the study of mitochondrial DNA variants of current populations showed the presence of a Paleolithic substrate attested by haplogroup U6. Specific North African populations: haplogroup U6 is dated to 47,000 ± 18,000 years (Rando et al, 1998, Plaza et al 2003..). The genetic structure of the North African populations is a mosaic due to the presence, next to 2 Mitochondrial diversity of the population Taforalt (12,000 years BP - Morocco): A genetic approach to the study of the settlement of North Africa U6, Eurasian haplogroups (such as H, U, K, J and T) and sub-Saharan haplogroups (L1, L2 and L3A) (Rando et al. 1998 Brakez et al. 2001, Plaza et al. 2003 Fadhlaoui-Zid et al. 2004). The importance of the contribution of the sub-Saharan contribution in the genetic composition of north African populations follows a south-north gradient (Krings et al. 1999 Brakez et al. 2001). Tunisian, Algerian populations, and Moroccan Berber as well as non Berber comprise a majority of individuals bearers of European haplogroups (from 56.4% to 88.4%). Mauritanian and Sahrawis have nearly as many individuals belonging to European haplogroups that sub-Saharan Africans (48% and 36.7% of European haplogroups 44% and 43.3% of African haplogroups) while the Tuareg populations, although speaking a Berber language, largely made up of African haplogroups (86.4%). The decrease in the frequency of the South Sub-Saharan haplogroups to the north is also observed in populations of the Nile Valley (Krings et al. 1999). This profile is consistent with North African populations share an origin with the people of the Middle East and Europe (Brakez et al. 2001, Plaza et al. 2003) that subsequently incorporated sub-Saharan sequences by an influx trans-Saharan genes (Brakez et al. 2001). This sub-Saharan contribution was probably recent, current view (Rando et al. 1998 Brakez et al. 2001). We present here the results we have obtained in the genetic study of thirty specimens of the population ibéromaurusienne Taforalt (12,000 years) (Ferembach 1962). Determining the genetic diversity of the population Taforalt will know better the ancestral composition of the North African populations and to clarify the chronology of TABLE 1. The 31 specimens studied population Taforalt. The Roman numeral indicates the burial number, the numeral indicates the number of skeletal or bone. migration. This genetic study will also help to clarify the pattern of origins men of Mechta El-Arbi which is a subject of controversy. THE ARCHAEOLOGICAL SITE Taforalt The archaeological site of Taforalt is located in the mountains of blessed Snassen, 1 km from the village of Taforalt and 55 km northwest of Oujda in Morocco. This is a cave 30 meters of opening and 28 m deep from front to back (Roche 1953) (Figure 1a). Taforalt cave was excavated from 1951 to 1955 by Father Roche. The stratigraphic study of the cave revealed 10 ibéromaurusiens levels (Epipalaeolithic) above a level Aterian (Roche 1953). EQUIPMENT The ancient population Taforalt The anthropological study of the site (Ferembach 1962) has highlighted the existence of skeletal remains of 86 adults and 98 children, in good condition, divided into 40 burials of the cave. The burials dated to 12,000 years ago, were discovered in the ten Epipalaeolithic levels. The men were Taforalt type Mechta El-Arbi (Figure 1b) and ibéromaurusienne culture. specimens studied Table 1 includes the 31 specimens (fragments of long bones, femur, fibula) taken from the collection of Taforalt, preserved at the Institute of Human Palaeontology in Paris. The Roman numeral indicates the burial number. The nomenclature of bone specimens Taforalt is a true enigma. The Arabic numerals following the burial number (Taf V-5, V-Taf 7 Taf V-18 ...) they corresponded to different individuals in the same grave? or is it the numbers of the different bones belonging to the same individual? It's a real problem, because each grave contained at least two individuals (I burial contained 9 people, 8 people were exhumed from the grave ... V)
The 21 specimens tested enough to give a good idea of the characteristics of the population Taforalt. None of these specimens is not related to the current sub-Sudanese specimens (absence of haplogroup L, gathering the typical African haplotypes SSA). The hypothesis of an origin of Saharan ibéraumaurusiens is not supported by our results, showing rather a Mediterranean settlement in Northern Africa until at least 12,000 years. Our results would support work based on dental analysis, morphometric facial and post-cranial cranio showing discrepancies between ibéromaurusiennes populations and their contemporaries in Sudan (Bermudez de Castro 1991 Irish 2000). The presence of a component in the sub-Saharan African populations north current was due to subsequent migrations to 12,000 years BP. All sequences specimens Taforalt have haplotypes belonging to Eurasian haplogroups. Among the majority haplogroups, H is considered as originating in the Middle East, the emergence of this haplogroup dated to 35,000 years, while haplogroup JT, from the Middle East also would have an age equal to 50,000 years; Finally haplogroup U6 from Northern Africa is proposed as a contemporary of haplogroup JT. The presence of individuals carrying a mitochondrial DNA belonging to haplogroup U6 shows in Taforalt persistence to 12,000 years BP of an ancestral component of North African origin. This result confirms both the North African emerging U6 and a likely evolution in situ an ancestor population Taforalt. The presence of H and JT translate Paleolithic migration flows from the Middle East. However, haplogroup H may indicate a strictly European origin component in the population Taforalt resulting from a later migration flow. Among the current population of North Africa studied to date, the Berber population of northern Morocco has similarities with the ancient population Taforalt. Indeed, Morocco Northern Berber have the lowest of haplogroup Saharan kind of rate (3.2%). In addition, all haplogroups observed in Taforalt are found in the current population, even haplogroup J / T. The latter, seldom represented in existing populations, is absent in the North African populations with the exception of the Berber population of northern Morocco (1.6%). These observations suggest a genetic continuity between ibéromaurusienne Taforalt population and current population occupying the same geographical area: the Berber population of northern Morocco. Our results reinforce anthropological studies that suggest that human type Mechta El-Arbi has not disappeared completely. The méchtoïdes characters are found in 8% of proto-historic and Punic tombs, in 3% of current populations of the Maghreb and highest percentage among the population of the Canary Islands (Ferembach 1962 Camps 1989). Our results give weight to dental studies, lithic and craniometric that suggest a regional continuity in North Africa since the Pleistocene terminal (Lubell et al. 1984 Irish 2000). CONCLUSION We have discussed in this article an innovative theme which is the study of the genetic diversity of the population Epipalaeolithic Taforalt. The hypothesis of a sub-Sudanese origin of ibéromaurusiens is rejected. A local North African origin is more likely. The study of polymorphisms of mitochondrial DNA from other ancient populations such as ibéromaurusienne Afalou population (Algeria) and Capsian populations still clarify our understanding of the peopling of North Africa.
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continue to deflect, distract, detract and mislead.
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Phylogeny and genetic structure of Tunisians and their position within Mediterranean populations
The mtDNA genetic pool of Tunisian populations contains three components: Eurasian lineages (H, HV, U (excepted U6), T, J, V, K, N, R, U5, I, W, X), Sub-Saharan lineages (L0, L1, L2, L3) and North African lineages (U6 and M1).
Among the Eurasian lineages, haplogroup H was the most frequent in Tunisia (Table 3, Supplementary material Appendix D). The frequencies of this haplogroup are extremely different even between close localities. For example, Bou Saaˆd showed the lowest value (5%) whereas Bou Omrane showed the highest value (47%) of haplogoup H and both of them belonged to the same governorate. In the second position, haplogroups T and U (except U5 and U6) are found in the majority of analyzed Tunisian samples.
Results from this study show that Tunisia is characterized by a mosaic genetic structure highlighted by the presence of a wide variety of haplogroups. 19 different haplogroups with several subclades were identified. Haplogroups were classified into three components: Eurasian, Sub-Saharan and North African lineages. Eurasian haplogroups are predominant followed by Sub-Saharan L lineages and specific North African clades: U6 and M1.
Our results are in accordance with previous ethnic studies using uniparental markers (mtDNA and Y chromosome) (Cherni et al., 2005, 2009; Ennafaa et al., 2011; Loueslati et al., 2006) and autosomal markers (Ancestry Informative SNP panel) (Khodjet-el-Khil et al., 2011).
The haplogroup U6 in Tunisia was likely diluted by migrations mainly from Near East installing the major haplogroups H, U and T. Other less frequent Eurasian lineages in Tunisia (HV, J, K and N), might also reflect the expansion of populations from the Near East. The presence of haplogroups (H1, H3, V, and U5b1) in the mtDNA genetic pool of the Tunisian population highlights a southward expansion from Europe (Fadhlaoui-Zid et al., 2011, Rhouda et al., 2009) in the Epi-Paleolithic period. Finally, two types of migrations seemed to explain the Sub- Saharan lineages in Tunisia; one from Eastern Africa (L0a1, L2a, L3e, L3f, L3h, L4, and L5), during pastoral movement (Ottoni et al., 2009) and the other from Western Africa (L1b, L2b, L2c, L2d, L2e, L3b, and L3d) (Richards et al., 2000; Salas et al., 2002). The presence of haplogroup L1c in Tunisia suggests a Central African origin (Salas et al., 2002). The introduction of Sub- Saharan lineages in Tunisia could be explained by the trans- Saharan trade of gold, salt and slaves.
quote: Haplogroup L1b roots deeply in the human mtDNA phylogeny and has the characteristic motif 16126, 16187, 16189, 16223, 16264, 16270, 116278, 16311.
[...]
Our results also point to a less ancient western African gene flow to Tunisia involving haplogroups L2a and L3b. Thus the sub-Saharan contribution to northern Africa starting from the east would have taken place before the Neolithic. The western African contribution to North Africa should have occurred before the Sahara’s formation (15,000 BP).
[...]
The dates for subhaplogroups H1 and H3 (13,000 and 10,000 years, respectively) in Iberian and North African populations allow for this possibility. Kefi et al.’s (2005) [data on ancient DNA could be viewed as being in agreement with such a presence in North Africa in ancient times (about 15,000–6,000 years ago) and with the fact that the North African populations are considered by most scholars as having their closest relations with European and Asian populations (Cherni et al. 2008; Ennafaa et al. 2009; Kefi et al. 2005; Rando et al. 1998). How- ever, considering the general understanding nowadays that human settlement of the rest of the world emerged from eastern northern Africa less than 50,000 years ago, a better explanation of these haplogroups might be that their frequencies re- flect the original modern human population of these parts of Africa as much as or more than intrusions from outside the continent. The ways that gene frequencies may increase or decrease based on adaptive selection, gene flow, and/or social processes is under study and would benefit from the results of studies on autosomal and Y-chromosome markers.
[...]
results reveal that Berber speakers have a foundational biogeographic root in Africa and that deep African lineages have continued to evolve in supra-Saharan Africa.
--Frigi et al.
quote: Focusing on the North African component at k = 6, we found that a migration event from North Africa to Europe would have occurred at least 6–10 generations ago (∼240–300 ya) in Spain, and at least 5–7 generations ago in France and Italy (Fig. 4).
[...]
Whereas inferred IBD sharing does not indicate directionality, the North African samples that have highest IBD sharing with Iberian populations also tend to have the lowest proportion of the European cluster in ADMIXTURE (Fig. 1), e.g., Saharawi, Tunisian Berbers, and South Moroccans. For example, the Andalucians share many IBD segments with the Tunisians (Fig. 3), who present extremely minimal levels of European ancestry. This suggests that gene flow occurred from Africa to Europe rather than the other way around.
--Laura R. Botiguéa,1, Brenna M. Henn et al
Gene flow from North Africa contributes to differential human genetic diversity in southern Europe (July 16, 2013)
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The Mesolithic–Neolithic transition in southern Iberia
Resumen: New data and a review of historiographic information from Neolithic sites of the Malaga and Algarve coasts (southern Iberian Peninsula) and from the Maghreb (North Africa) reveal the existence of a Neolithic settlement at least from 7.5 cal ka BP. The agricultural and pastoralist food producing economy of that population rapidly replaced the coastal economies of the Mesolithic populations. The timing of this population and economic turnover coincided withmajor changes in the continental and marine ecosystems, including upwelling intensity, sea-level changes and increased aridity in the Sahara and along the Iberian coast. These changes likely impacted the subsistence strategies of the Mesolithic populations along the Iberian seascapes and resulted in abandonments manifested as sedimentary hiatuses in some areas during the Mesolithic–Neolithic transition. The rapid expansion and area of dispersal of the early Neolithic traits suggest the use of marine technology. Different evidences for a Maghrebian origin for the first colonists have been summarized.
The recognition of an early North-African Neolithic influence in Southern Iberia and the Maghreb is vital for understanding the appearance and development of the Neolithic in Western Europe. Our review suggests links between climate change, resource allocation, and population turnover.
Cortés-Sánchez, Miguel Et al.
The Mesolithic–Neolithic transition in southern Iberia
quote:Haplogroup H dominates present-day Western European mitochondrial DNA variability (>40%), yet was less common (~19%) among Early Neolithic farmers (~5450 BC) and virtually absent in Mesolithic hunter-gatherers. Here we investigate this major component of the maternal population history of modern Europeans and sequence 39 complete haplogroup H mitochondrial genomes from ancient human remains. We then compare this 'real-time' genetic data with cultural changes taking place between the Early Neolithic (~5450 BC) and Bronze Age (~2200 BC) in Central Europe. Our results reveal that the current diversity and distribution of haplogroup H were largely established by the Mid Neolithic (~4000 BC), but with substantial genetic contributions from subsequent pan-European cultures such as the Bell Beakers expanding out of Iberia in the Late Neolithic (~2800 BC). Dated haplogroup H genomes allow us to reconstruct the recent evolutionary history of haplogroup H and reveal a mutation rate 45% higher than current estimates for human mitochondria.
--Brotherton P1, Haak W, Templeton J,
Nat Commun. 2013;4:1764. doi: 10.1038/ncomms2656.
Neolithic mitochondrial haplogroup H genomes and the genetic origins of Europeans.
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