The Complex and Diversified Mitochondrial Gene Pool of Berber Populations
C. Coudray1∗ , A. Olivieri2, A. Achilli2,3, M. Pala2, M. Melhaoui4, M. Cherkaoui5, F. El-Chennawi6, M. Kossmann7, A. Torroni2 and J. M. Dugoujon1 1Laboratoire d’Anthropobiologie, CNRS FRE2960, Universite´ Paul Sabatier, Toulouse, France
Summary The mitochondrial DNA variation of 295 Berber-speakers from Morocco (Asni, Bouhria and Figuig) and the Egyptian oasis of Siwa was evaluated by sequencing a portion of the control region (including HVS-I and part of HVS-II) and surveying haplogroup-specific coding region markers. Our findings show that the Berber mitochondrial pool is characterized by an overall high frequency of Western Eurasian haplogroups, a somehow lower frequency of sub-Saharan L lineages, and a significant (but differential) presence of North African haplogroups U6 and M1, thus occupying an intermediate position between European and sub-Saharan populations in PCA analysis. A clear and significant genetic differentiation between the Berbers from Maghreb and Egyptian Berbers was also observed. The first are related to European populations as shown by haplogroup H1 and V frequencies, whereas the latter share more affinities with East African and Nile Valley populations as indicated by the high frequency of M1 and the presence of L0a1, L3i, L4∗, and L4b2 lineages. Moreover, haplogroup U6 was not observed in Siwa. We conclude that the origins and maternal diversity of Berber populations are old and complex, and these communities bear genetic characteristics resulting from various events of gene flow with surrounding and migrating populations.
Introduction Even if archaeological and paleoanthropological records tes- tify to the ancient (Paleolithic) human occupation of North Africa, the evolution of human groups living in that area is still unclear. This is mainly due to the large number of suc- cessive prehistoric and historic events that occurred in that area after the arrival of the first modern humans. In North Africa, the presence of Berbers – a term to denote those populations which speak a Berber language (Camps 1980) – is well described since the Capsian (10,000–4700 years ago),
although this industry derived from oldest cultures. Never- theless, it was only during the Neolithic transition (around 6000 years ago in the Saharan areas and 5000 years ago in the Maghreb) that North Africa was incontestably marked by various cultural events. Then, Berbers experienced a long and complicated history with many invasions, conquests and migrations by Phoenicians, Romans, Vandals and Byzantines (Brett & Fentress 1996). The most significant event was the Arab conquest, begun during the 7th century, when North Africans were converted to Islam, and Arabic became the of- ficial unique language employed. In spite of strong resistance, Berbers acquiesced to Arab authority. Refractory groups were driven out and constrained to more isolated areas. This trou- bled past directly influenced the geographical distribution of Berber communities which are nowadays scattered in a vast region extending from Mauritania to Egypt (Siwa oasis) and from the Sahara desert to the Moroccan Atlas mountainous
areas. Over the course of time, the various populations that migrated to North Africa have probably left a footprint in the gene pool of modern Berbers. A detailed analysis of human mtDNA could be a power- ful tool in reconstructing the population history of a region, such as Northern Africa, which is at the crossroads of Europe, the Middle East, and sub-Saharan Africa, regions for which mtDNA phylogeography is known in detail. Many studies have attempted to describe the genetic diversity of Berber populations, evaluating mitochondrial DNA (mtDNA) se- quence variation (Achilli et al. 2005; Brakez et al. 2001; Cherni et al. 2005; Fadhlaoui-Zid et al. 2004; Krings et al. 1999; Loueslati et al. 2006; Macaulay et al. 1999; Olivieri et al. 2006; Plaza et al. 2003; Rando et al. 1998; Stevanovitch et al. 2004). In this study, we present sequence data from mtDNA, detected by surveying a portion of the control re- gion, including hypervariable segment I (HVS-I) and most of hypervariable segment II (HVS-II) and haplogroup diag- nostic RFLP markers. We studied four Berber populations: three from Morocco (Asni, Bouhria and Figuig) and one from Egypt (Siwa oasis). Our main objectives were to describe the maternal lineages of the Berbers, to determine the genetic contributions to their mitochondrial pool from surrounding populations (particularly from the sub-Saharan area, south- ern Europe, and the Near East), and to link their present-day genetic diversity to the numerous human migrations which affected North Africa. Materials and Methods Populations We studied 295 unrelated and healthy individuals from four dif- ferent Berber populations. Three groups were from Morocco, Asni (N = 53), Bouhria (N = 70) and Figuig (N = 94), and one was from the Egyptian oasis of Siwa (N = 78). The geograph- ical location of the sampled sites is presented in Figure 1. Asni is located in the Tacheddirt valley in the High Atlas Mountains (47 km from Marrakech). The Berbers from Asni speak Chleuh
and belong to the Rhiraya tribe. Sidi Bouhria is located in north- eastern Morocco (Oujda wilaya). Berbers from Bouhria belong to the Beni Moussi Roua fraction of the Beni Iznasen tribe. Figuig is located in eastern Morocco, at the Algerian border, at the juncture of the High Plateaus and the north-western edge of the Sahara. It is an oasis encompassing seven villages, and the Berber population that we studied lives in the largest of these, Zenaga. The Siwa oasis is located in the western Egyptian desert, at 300 km from the Mediterranean coast and 25 km from the Libyan border. This population speaks Siwi, a Berber language. Before blood collection, people were interviewed in order to ascertain their ethnic origins and to obtain informed consent. All individuals and their families included in this study have been living in the area of interest for at least three generations. In addition, mtDNA haplogroup information from 58 European, Near Eastern, and African populations was used for comparative analyses (see Table 1). DNA Extraction For the Moroccan Berbers, genomic DNA was isolated from blood samples using a standard proteinase-K digestion fol- lowed by phenol-chloroform extraction and ethanol precipita- tion, while for the sample from Siwa, genomic DNA was iso- lated from blood using the NucleoSpin Blood Mini Kit (BD Biosciences Clontech, San Jose, USA). MtDNA Amplification, Sequencing and RFLP Screening A portion of the mtDNA control region encompassing the entire HVS-I and part of HVS-II was PCR amplified using primer pairs F15877 (5-CAAATGGGCCTGTCCTTGTA-3) and R468 (5-GGAGTGGGAGGGGAAAATAA-3) for sam- ples from Bouhria, Asni and Siwa, and primers F15973 (5- AACTCCACCATTAGCACCCA-3) and R296 (5-GGAAAT TTTTTGTTATGATGTCT-3) for the Figuig sample. DNA purification was undertaken using the QIAquick PCR Purifica- tion Kit (QIAgen, Courtaboeuf, France). Reactions were carried out using the BigDye Terminator (version 3.1) Ready Reaction Cycle Sequencing kit, with AmpliTaq DNA polymerase (PE, Applied Biosystems, Foster City, USA). Sequences were run in
an automatic Sequencer ABI PRISM 3730 (PE, Applied Biosys- tems). Both strands were sequenced using primers F15973 and R263 (5 -TGGCTGTGCAGACATTCAAT-3 ), and only con- firmed deviations from the reference sequence were considered. In addition to the control region sequencing, and in order to clas- sify mtDNAs unambiguously within haplogroups, coding region diagnostic markers were surveyed (see supplementary material table). Statistical and Phylogenetic Analyses The mtDNA sequences were aligned using the BIOEDIT soft- ware (Tom Hall, Carlsbad, USA). Only segments from positions 16033 to 220 according to the revised Cambridge Reference Sequence rCRS (Anderson et al. 1981; Andrews et al. 1999) were considered for analysis. Haplogroup assignment followed the basic classification scheme of Torroni et al. (1996; 2006). Additional reports (Achilli et al. 2004; Achilli et al. 2005; Behar et al. 2008; Kivisild et al. 2004; Macaulay et al. 1999; Olivieri et al. 2006; Quintana-Murci et al. 2004; Quintana-Murci et al. 1999; Richards et al. 2000; Richards et al. 1998; Salas et al. 2004; Salas et al. 2002; Shen et al. 2004) were consulted to fur- ther delineate the haplogroups and to assign geographical labels (West Eurasian, East Eurasian, North African, and sub-Saharan African). Population internal genetic diversity parameters (num- ber of different sequences, haplotype diversity, nucleotide diver- sity, and mean number of pairwise differences) were calculated for each sample using Arlequin 3.0 software (Excoffier et al. 2005).
Based on the available haplotypes, population-specific median networks were generated with the median joining algorithm (Bandelt et al. 1995; 1999) using the NETWORK 4.5 software program developed by Fluxus Technology Limited (Sudbury, UK http://www.fluxus-engineering.com). The weighting scheme for the nucleotide positions used in this analysis (nps 16033–220) followed Richards et al. (1998). Subsequently, the most parsi- monious tree was inferred and drawn manually and confirmed by using the Network MP-calculation option. For summarizing population relationships, Principal Component Analysis (PCA) of haplogroup frequencies was performed using the Excel Stat package (Microsoft). PCA provides a method for representing frequency data in a Euclidian space so that the results can be visually examined for structure (Greenacre 1992). For the PC analyses, we used the frequencies of the following haplogroups: H (including H∗ and all H subhaplogroups), HV0 (including HV0∗ and V), HV (including HV∗ and HV1), R0 (represented mainly by R0a), J, T, U (including all U lineages except U6 and K), U6, K, N1 (including N1 and I lineages), N2 (represented mainly by W), X, M (including M∗ and all M subhaplogroups except M1), M1, L0, L1, L2, L3, and L4-L5. Population ge- netic structure was tested by analysis of variance (AMOVA) and F statistics (Excoffier et al. 1992; Weir & Hill 2002) using the Arlequin program (Excoffier et al. 2005). The significance of the covariance components associated with the different levels of genetic structure was tested using a non-parametric permu- tation procedure (Excoffier, Smouse & Quattro 1992). For this analysis, population samples were grouped into major geographic
areas: North Africa, East Africa, sub-Sahara, the Near East and Europe. We estimated the genetic variation among populations within the same geographical group (FSC), among geographical groups (FCT), and among populations across the entire study area (FST). Molecular pairwise FST indexes between populations were calculated from haplogroup frequencies, and their signifi- cance was evaluated with a nonparametric permutation test using the Arlequin package (Excoffier, Laval & Schneider 2005). Results MtDNA Sequence Diversity in the Four Berber Populations The total number of different control region haplotypes in our sample of 295 Berbers was 127. Among these, only 9 haplotypes were shared between two or three populations (4 between Asni and Bouhria, 3 between Asni and Figuig, 1 between Bouhria and Figuig and 1 among Asni, Bouhria and Figuig). Among the 53 Berbers from Asni, 42 different se- quences belonging to 19 distinct haplogroups were found. The 70 Berbers from Bouhria were characterized by 38 dif- ferent sequences and 26 haplogroups/sub-haplogroups. In the 94 Berbers from Figuig, 32 different sequences and 20 haplogroups were detected. Finally, among the 78 Egyptian Berbers, the 25 different observed sequences were subdivided into 19 distinct haplogroups. Haplogroup frequencies in the four Berber populations are reported in Table 2. The four mtDNA pools show different compositions but overall harbor a high frequency of Eurasian haplogroups (H, HV0, HV, R0, J, T, U w/o U6, K, N1, N2, and X), and a relatively lower frequency of sub-Saharan L lineages (L0, L1, L2, L3, and L4-L5). Both North African- specific clades U6 and M1 are represented but with extremely different distributions. In detail, Asni and Bouhria samples showed slightly similar genetic profiles whereas those from Figuig and Siwa were quite different, in particular because of their higher frequency of L and M1 haplogroups, respec- tively. When lineages are considered one by one, several dif- ferences appear within Moroccan populations, and between Moroccans and Egyptians. For example, among West Eurasian lineages, haplogroup H was the most common in Morocco whereas haplogroups U (without U6 and K) and K were the most represented in Siwa. As for the sub-Saharan lineages in Morocco, the Berbers from Figuig were found to harbor the highest proportion of L mtDNAs. M1 mtDNAs were instead much more frequent in Siwa than in Morocco, whereas U6 was only observed in Morocco with the highest frequency in Asni. Table 3 shows the descriptive parameters of the analyzed populations. Gene diversity values in Asni and Bouhria sam- ples are very similar whereas Figuig and Siwa samples show a reduced diversity level, Siwa showing the lowest values. Forthe nucleotide diversity and mean number of pairwise differ- ences, the sample from Figuig tends to have slightly higher values. The observed reduced diversity in the maternal gene pool of the Berbers from Figuig and Siwa could be attributable to genetic drift because of the relative isolation of the oases where they live. Genetic diversity values were compared with those of a set of North African Berber populations. The diversity indices obtained here for Asni (0.983), Bouhria (0.966) and Figuig (0.956) samples are in full accordance with previous data. The observed high level of mtDNA sequence diversity is in the range of values generally found in North Africa: 0.941 for the Souss Valley (Brakez et al. 2001), 0.942 for Moz- abites (Corte-Real et al. 1996; Macaulay et al. 1999), 0.939, 0.975 and 0.964 for Chenini-Douiret, Sened and Matmata (Fadhlaoui-Zid et al. 2004), respectively. Within all Berber populations, the Siwi show the lowest value of sequence di- versity (0.921). The Phylogeography of North African Berbers To visualize the mtDNA diversity of the four Berber sam- ples, a tree was constructed and presented in Figure 2 (for mitochondrial sequences, see additional file 1). Eurasian Lineages in Moroccan and Egyptian Berbers Thirty-three, 57, 47 and 45 individuals out of 53, 70, 94 and 78 samples from Asni, Bouhria, Figuig and Siwa (respec- tively) carried mtDNAs that are typical of western Eurasian populations, particularly from Europe. Haplogroup H is by far the most frequent in western Eu- ropean populations. It accounts for 40%-50% of the mtDNA pool in most of Europe, and 20%–30% in the Near East and the Caucasus region (Richards et al. 2000; Roostalu et al. 2007). It is thought to have evolved in the Near East and Mid- dle East ∼23,000–28,000 years ago and to be involved in the late glacial expansions that took place into Europe ∼13,000– 15,000 years ago, mainly from the Iberian refuge (Achilli et al. 2004; Loogvali et al. 2004; Pereira et al. 2005; Torroni et al. 1998). H is observed at high frequencies in Moroc- can samples (37.7%, 37.1% and 24.4% in Asni, Bouhria and Figuig, respectively) whereas it accounts for only 1.3% of the Egyptian Berbers. Different subclades of H lineages are present in Asni (H∗, H1, H3, and H7), Bouhria (H∗, H1, H2, H3, H5, H6, and H11), Figuig (H1 and H8), and Siwa samples (H1). Among these, H1, which is the most common sub-clade of H in Europe (Achilli et al. 2004; Finnila et al. 2001; Herrnstadt et al. 2002; Loogvali et al. 2004), is also the most represented in the Berber samples. HV0 probably originated in Europe before the Last Glacial Maximum. MtDNAs belonging to this haplogroup are gen- erally rare and apparently scattered throughout Europe,although they are less uncommon in the Mediterranean area (including northwest Africa) (Torroni et al. 2001). Indeed, HV0 mtDNAs were observed in 6 subjects from Asni, 10 from Bouhria, 8 from Figuig and 10 from Siwa. HV0 lineages were represented by both HV0∗ (2, 9, 5 and 10 sequences in Asni, Bouhria, Figuig, and Siwa, respectively) and V sub-clades. Vsequences generally represent 5% of European mtDNAs and tend to be restricted to western, central, and northern Europe (Torroni et al. 1998; Torroni et al. 2001). Four, one and three individuals belonging to this haplogroup were detected in the Asni, Bouhria and Figuig samples, respectively, whereas V was not observed in Siwa.
Only one individual from Bouhria showed a HV1 sequence (a sister clade of HV0 and H) and one Egyptian Berber bore a HV∗ sequence. Haplogroup R0a, a sister clade of HV, oc- curs in populations from the Near East, the Caucasus, and Mediterranean Europe. It was found in two Berbers from Asni and two Siwi. J and T sister clades might have originated in the Near East ∼50,000 years ago, but they could have been re- cently introduced into Europe during the Neolithic ∼10,000 years ago (Finnila et al. 2001; Palanichamy et al. 2004; Richards et al. 2000). Haplogroup J is observed in popula- tions from the southern Caucasus, the Near East, and NorthAfrica (Brakez et al. 2001; Maca-Meyer et al. 2001; Plaza et al. 2003; Richards et al. 2000). It is not found in the Asni and Bouhria samples, whereas the Berbers from Figuig harbor five J1c, three J2, and three J2a mtDNAs, and the Berbers from Siwa show four J2 mtDNAs. The most numerous T sequences were observed in Bouhria (six T1a and one T2b) while the Figuig samples exhibited three T2 and the Siwa sample only one T1a mtDNA. Haplogroup T was not detected in Asni. Eurasian U macrohaplogroup is subdivided into U1 to U9 (with the exception of U6 restricted to North Africa) and K lineages. U has an extremely broad geographical distribution and accounts for about 20% of European mtDNA sequences (Herrnstadt et al. 2002). In this study, it is represented by four haplogroups: U2, U3, U4 and U5. U2 can be found at low frequencies in populations of western Asia and the Caucasus. Here, it was observed only in the Berbers from Bouhria, by two subclades, U2b (1 sequence) and U2e (2 sequences). U3 is observed at 1.1% (U3a) and 1.3% (U3) in samples from Figuig and Siwa, respectively. This haplogroup has a frequency peak in the Near East (Achilli et al. 2007). U4 haplogroup is spread at moderate frequencies all over Europe, western Siberia, and southwestern Asia (Richards et al. 2000). Only one sequence from Bouhria belongs to U4. U5, one of the most ancient subhaplogroup of U, occurs in most cases as occasional haplotypes that are derived from European lineages (Achilli et al. 2005; Richards et al. 2000). In 16.7% of Siwi samples, it appears as U5b, a lineage suggesting back-migration of people from Europe to the South (Torroni et al. 2006). One Berber from Asni also bears this sequence. K lineages are observed at 3.8%, 5.7% and 11.5% in samples from Asni, Bouhria and Siwa, respectively. N1b1 is a minor mtDNA haplogroup that has been ob- served at marginal frequencies in European and Near Eastern populations (Palanichamy et al. 2004; Richards et al. 2000). It occurs at a very low frequency in both Asni (1.9%) and Bouhria (1.4%) samples. N2 sequences, represented by W∗ haplogroup, is only present in two Berbers from Bouhria. X∗ mtDNAs occur at low frequency in Asni (1.9%), Bouhria (2.9%), Figuig (1.1%) and Siwa (3.8%) samples. All major European mtDNA haplogroups are most likely of Middle Eastern origin (Torroni et al. 1996, 2001). Their presence at high frequencies in Berber samples indicates ei- ther gene flow from Europe to North Africa and/or a prob- able common ancestry of populations of both shores of the Mediterranean Sea. Despite total West Eurasian lineages oc- cur at equally high frequency in the four Berber samples, they exhibit a different distribution between Morocco and Egypt, with the Siwi showing the lowest frequency of the West European H but the highest frequencies of HV0∗ and K haplogroups which are more prevalent in East Eurasia and the Near East.Sub-Saharan Llineages in Moroccan and Egyptian Berbers L African lineages which account for 22.6%, 12.9%, 44.7% and 24.4% of Asni, Bouhria, Figuig and Siwa mitochon- drial pool are represented by six haplogroups: L0, L1, L2, L3, L4 and L5 (Gonder et al. 2007; Torroni et al. 2006). Haplogroup L0 is the earliest offshoot of the mtDNA tree in Africa that appears as a sister group to the branch that holds all other haplogroups (Salas et al. 2002). L0a subclade prob- ably originated in Eastern Africa and is common in Eastern, Central, and South Eastern Africa. One L0a1 sequence was found in an Egyptian Berber. Although L0f subclade is rare and geographically confined to eastern Africa, it was observed in four Berbers from Figuig. Haplogroup L1 appears in our dataset as L1b and L1b1 sub-clusters. L1b and L1b1 are particularly concentrated in Western Africa, although with some overflow into Central and North Africa (Salas et al. 2002). L1b is observed in 2 sequences from Bouhria and L1b1 in 4, 3, 6, and 1 individuals from Asni, Bouhria, Figuig, and Siwa, respectively. Three sequences from Asni, two from Bouhria, four from Figuig and four from Siwa, belong to L2 lineages. They appear as four clusters, L2a (in Bouhria and Siwa), L2a1 (only in Moroccan populations), L2b1 (in Asni), and L2d2 (in Figuig). While L2a is the most common and widespread L2 subclade all over Africa, L2b is largely restricted to western Africa and L2d is rather rare but also appears to be restricted to Western Africa (Salas et al. 2002; Torroni et al. 2001). L3 lineages appear in our dataset as three clades, L3b, L3e and L3i (Salas et al. 2002). L3b is widely spread in West- ern African populations and Bantu-speaking South Eastern Africans and it is also observed in Ethiopia (Kivisild et al. 2004; Salas et al. 2002). L3b accounts for 8.5% of Figuig mtDNA sequences while it is absent in the three other pop- ulations. L3e is prevalent in central (∼20%) and western (∼11%) Africa but it is less abundant (∼3%) in eastern Africa. It orig- inated from the Sudan (Bandelt et al. 2001). It is observed as 9 L3e1 in Siwa, as 4 L3e2 in Figuig, and as 4, 2 and 16 L3e5 in Asni, Bouhria and Figuig samples, respectively. L3e1 is the oldest and most diverse clade among L3e types. Albeit prac- tically omnipresent in sub-Saharan and South Eastern Africa, it is supposed to have had a western/central African origin. L3e2 is omnipresent in Africa (Bandelt et al. 2001). L3e5 is considered to be restricted to North Africans (Fadhlaoui-Zid et al. 2004; Torroni et al. 2006). L3i haplogroup seems to be restricted to Ethiopia and Yemen (Kivisild et al. 2004). One individual from the Siwa oasis exhibits an L3i2 sequence (Behar et al. 2008). L4 haplogroup, which is an early branch of L3 lineages, is only represented in Berbers from Siwa by 2 L4∗ (L4 subclade not defined) and 1 L4b2 sequences (Behar et al. 2008). L4 isan east African lineage and reveals high sequence diversity in Ethiopians (Kivisild et al. 2004). L5 has been observed at low frequency only in eastern Africa (Kivisild et al. 2004). It is present in one individual from Asni. L lineages are specific of the mitochondrial pool of sub- Saharan populations. Their presence at relatively high fre- quencies in North African Berbers could be attributed to a gene flow from the Saharan areas. However, these ex- changes do not seem to have the same geographical origin: in Morocco, frequencies of L1b, L2b, L2d, L3b and L3e5 haplogroups indicate a predominantly west sub-Saharan in- fluence, whereas in Egypt, L0a, L3e1, L3i and L4 lineages underline migration routes from eastern Africa. M Asian, M1 and U6 North African Lineages in Moroccan and Egyptian Berbers M1 is a subclade of southwest Asian ancestry that moved to Africa about 40,000 to 45,000 years ago (Olivieri et al. 2006). Its highest frequencies are found in Egypt (∼18%) and in eastern Africa (∼11% in Ethiopia and Somalia). It is not uncommon in the Mediterranean basin showing a peak in the Iberian Peninsula and it is also observed in the Middle East (Gonzalez et al. 2007). In our dataset, M1 is represented by two subclades, M1a, particularly abundant in Ethiopia, and M1b detected only in the Mediterranean area (Olivieri et al. 2006). We observed two M1a, three M1b, two M1a1, and 13 M1a1 sequences from Asni, Bouhria, Figuig, and Siwa samples, respectively. Finally, one mtDNA from Siwa was found to belong to the South Western Asian M33a clade (Abu-Amero et al. 2008). U6 is a North African clade, that similar to M1, is of south- west Asian ancestry and most likely entered Africa ∼40,000– 45,000 years ago (Olivieri et al. 2006). U6 is distributed from the Near East to Northwest Africa where it is found at its highest frequencies (Maca-Meyer et al. 2003; Olivieri et al. 2006). It was also observed at a sparse distribution in popu- lations from the southwestern part of the Iberian Peninsula (Plaza et al. 2003). In our dataset, U6 as a whole is only ob- served in the Moroccan samples, where it is represented by its most significant subgroup, U6a (five mtDNAs in Asni, one in Bouhria, and three in Figuig). In Asni, one U6d mtDNA was also observed. Although U6 is considered specific of Berber populations, its absence in the Siwa sample suggests a differentiation be- tween Berber-speakers living at the extremes of their geo- graphical distribution range. An opposite trend is that ob- served for M1 that is much more common in the Berbers from Egypt than in the Berbers from Morocco. This reveals a more significant east African influence in the North Eastern African genetic pool than in the North Western one.Mitochondrial Diversity in a Broad Geographical Context To observe the mtDNA patterns of Berber populations within a broad geographical context, we performed a first Principal Component Analysis (PCA) including data from 62 Euro- pean, Near Eastern and African populations (Fig. 3a). Princi- pal components 1 and 2 (axes 1 and 2 in Fig. 3) account for 46% of the total variation (32% and 14%, respectively). The two-dimensional pattern displays distinct population clusters: the sub-Saharan populations located in the most positive por- tion of axis 1; the East Africans and the Near Eastern groups in the upper-right and upper-left quadrant, respectively; the Europeans and some East Mediterranean populations located in the negative portion of axis 1; and North Africans sand- wiched between the sub-Saharan cluster and European pop- ulations, but closer to the latter. If we detail the position of our Berber samples in the overall mitochondrial diversity, the Moroccan Berbers from Bouhria are closest related to Euro- peans whereas Figuig are furthest away, with Asni being in an intermediate position. Siwi Berbers are far from the other three Moroccan samples and have closest relationships with East Mediterranean populations of Jordania, Syria, Israel and Iraq. The contribution of each haplogroup to the first two Prin- cipal Components is illustrated in Figure 3b. The population positions on the PCA are explained by the distinction between African and Eurasian haplogroups. As expected, sub-Saharan samples are characterized by high frequencies of L lineages (L0, L1, L2 and L3) whereas Europeans are mainly associ- ated with lineages H, HV0, U, K, T, N2 and X. Then, East Africans show high frequencies of M1 and L4-L5 lineages, and Near Easterners are mainly associated with lineages HV, R0, J, N2, M, and N1. Finally, the centered position of North Africans is explained by higher frequencies of West Eurasian lineages with respect to the African haplogroups, by the small frequencies of East Eurasian and M1 lineages, and by the presence of U6 haplogroup. Overall, the PCA plot based on mitochondrial haplogroup frequencies (Fig. 3a) reflects a clear differentiation between Berbers and sub-Saharans but a close relationship between Berbers and South Western European populations. This fits within the phylogeographical patterns established by previous genetic research (Plaza et al. 2003; Rando et al. 1998; Rosa et al. 2004). We carried out analyses of variance (AMOVA) on the pop- ulations used for the PC analysis (Table 4). For this, population samples were grouped into major geographical areas: North Africa, sub-Sahara, East Africa, the Near East and Europe. Results show that most of the genetic variation (83.38%) occurs within the populations whereas there is a relatively little (2.61%) but significant differentiation among popula- tions within the same geographical region (FSC = 0.030). A14.01% of the variance was attributed to differences among geographical groups (FCT = 0.140). This means that popu- lations within each group are relatively homogeneous, that groups are significantly differentiated and that the mitochon- drial diversity is structured according to a geographical classi- fication. Moreover, FST index is definitely higher than 0.005 (FST = 0.166) showing a strong diversification among popu- lations. By performing AMOVA between pairs of geographi- cal groups, we measured the population structuring observed in the PC analysis (Table 4). We found that 12.53% of the variance was attributed to differences among North African and sub-Saharan groups, 4.08% among North Africans and East Africans, 1.39% among North Africans and Near East- ern groups, and 5.01% among North Africans and Europeans. However, the highest value is observed between Europeans and sub-Saharans (25.72% of the total variance). Berber Mitochondrial Diversity An additional PC analysis was performed only on the ten Berber populations of our database (Fig. 4a) by considering their haplogroup frequencies (Table 5). The first two principal components account for 41% of the total variation (22% and
19%, respectively). The first axis reveals a clear differentiation between Siwa samples and all other Berber populations from Morocco, Tunisia and Algeria, while the second axis shows the Berbers from Chenini-Douiret (Tunisia) as the following most differentiated group. Our Moroccan samples are located at the centre of the graph. Locations of the population on the PCA plot are explained by the distinction between M, L4-L5, M1, X, K, U, R0, HV0, and L0 haplogroups on positive values (by descending order of contribution on axis 1) and H, T, HV, L1, L2, U6, N1, J, L3, and N2 lineages for negative values of axis 1 (Fig. 4b). In particular, the Siwa mitochondrial pool is characterized by the highest frequencies of M1 lineages (16.7%, from 0% to 11.3% in the Maghreb, see Table 5), by the lowest frequencies of H haplogroups (1.3%, from 13.1% to 37.7% in the Maghreb) and by the absence of the U6 North African lineage (from 0% in Chenini-Douiret and Jerba to 28.3% in Mozabites) (Corte-Real et al. 1996; Loueslati et al. 2006; Macaulay et al. 1999). Comparison of FST values from pairs of populations re- veals that, with the exception of some pair-groups (Asni vs. Bouhria, Asni vs. Souss, Asni vs. Sened, Asni vs. Matmata, Bouhria vs. Souss, Bouhria vs. Matmata, Figuig vs. Matmata, Souss vs. Sened, Souss vs. Matmata, Souss vs. Jerba, Sened vs. Jerba), Berber samples displayed significant differences inbroad phylogeographic context, in particular through a com- parison with population groups from southern Europe, the Saharan areas and the Near East. Analysis of the 62 popula- tions included in our haplogroup frequency database revealed that Berber populations are genetically close to southern Europeans, but significantly differentiated from sub-Saharan groups. This peculiarity is explained by their mitochon- drial genetic structure, characterized by an overall high fre- quency of Western Eurasian haplogroups, a somehow lower frequency of sub-Saharan L lineages, and a significant (but differential) presence of North African haplogroups U6 and M1. The genetic proximity observed between the Berbers and southern Europeans reveals that these groups shared a com- mon ancestor. Two hypotheses are discussed: one would date these common origins in the Upper Paleolithic with the ex- pansion of anatomically modern humans, from the Near East to both shores of the Mediterranean Sea; the other supports the Near Eastern origin, but would rather date it from the Neolithic, around 10,000 years ago (Ammerman & Cavalli- Sforza 1973; Barbujani et al. 1994; Myles et al. 2005; Rando et al. 1998). Common polymorphisms (i.e. those defining H and V lineages) between Berbers and south Europeans also could have been introduced or supported by genetic flows through the Straits of Gibraltar. For example, genetic ex- changes could have taken place during prehistory, while Eu- ropean populations retreated from ice sheets and expanded from refuge, around 15,000 years ago (as evidenced by the H and U5b mitochondrial lineages). Alternatively, these ex- changes could have occurred during history, with the invasion and the occupation during nearly seven centuries (from the 8th to the 15th century) of the Iberian Peninsula by Almora- vide then Almohade Muslim Berber troops. The differentiation observed between North Africans and sub-Saharan populations shows, first, that settlement of these areas was achieved by different migration waves and, then, that a genetic diversity was already observable in Africa since very old times. However, the Berber genetic heritage con- sists of a relatively high frequency of L lineages from various parts of Africa (i.e. L0a, L3i, L4, and L5 clades are from East Africa, L1b, L2b, and L3b are from West Africa, and L3e originated in the Sudan). It poses a question about the Sa- hara desert role in population movements and exchanges. It should be specified that the Sahara was not always a desert, because it also underwent enormous variation between wet and dry, offering green spaces favorable for human occupation and animal domestication (Aumassip et al. 1994; Said & Faure 1990). Thus, this is plausible that exchanges between African prehistoric populations took place; exchanges during which markers typical of sub-Saharan groups would have been in- troduced into the Berber gene pool. Contacts between North Africa and great sub-Saharan empires (such as those of Ghana,of Mali, or the Songhai Empire) are also reported by history during trans-Saharan trade of gold, salt and slaves. The second analysis we conducted aimed to measure the relationship between the Berber communities on their whole geographical habitat. Although the Berber populations have the same overall mitochondrial genetic composition, the dis- tribution of some markers is not the same along the Berber- speaker habitats. Our results highlighted a clear genetic differ- entiation between Berbers from the Maghreb and Egyptian Berbers. The first seems to be more related to European populations as shown by haplogroup H1 and V frequencies, whereas the latter share more affinities with East African and Nile Valley populations as indicated by the high frequency of M1 and the presence of L0a1, L3i, L4∗, and L4b2 lin- eages. Moreover, haplogroup U6 was not observed in Siwa. Probably, such a maternal diversity between North African Berbers would have been the result of a conjunction of several geographical, prehistoric, and historic factors which guided contacts (and thus exchanges) between local populations and migrating groups. First, in addition to the geographical dis- tance, which certainly increases the genetic distance, the geo- graphical location of Berber populations is very peculiar: the Berbers from the Maghreb are at the end of a long migration route, whereas Berbers from Siwa are rather in a crossroads be- tween the Middle East, East Africa, sub-Saharan areas and the North African corridor. Therefore, meetings and exchanges between local and migrating populations were not identical in North West and North East Africa. In addition, prehistory and history of the populations from Maghreb are different from those of the Egyptian group. For the Siwa oasis, there is very little information on which exact prehistoric period was the starting point of Berber culture. However we know that throughout history, the oasis was crossed by successive human groups, like pilgrims traveling to Mecca, Mediter- ranean tradesmen, or Sahelian slave merchants. Siwa was also repopulated by Libyan Berber-speakers driven from their land by Arab conquerors. Lastly, it experienced a period of decline and has faced, between the ninth and twelfth centuries, a dras- tic demographic reduction of its population (Fakhry 1973). The current gene pool of the Siwa people would therefore be the result of these various genetic exchanges which occurred in the past. Overall, it is clear that neither the Strait of Gibraltar nor the Sahara desert appears to have forced the movements and exchanges between South Europeans, North Africans, and sub-Saharans. This interaction took place since prehistory in a human and genetic framework already diversified, and the maternal ancestors of the Berbers were able to exchange some markers with surrounding populations with different cultures and genetic pools. Influences from the Middle East and East Africa are marked in Siwa, while southwestern European in- fluences are observed in the Maghreb. Although the origin of these Eurasian and sub-Saharan lineages in the mitochondrial pool of Berbers is still questionable (whether due to common ancestry or past and/or current gene flow), certainly they were not diluted by the many historical invasions and migra- tions, leaving a clear maternal footprint in the contemporary populations. We conclude that the origins and diversity of Berber pop- ulations are old and complex, and these communities bear genetic characteristics resulting from various events of gene flow with surrounding and migrating populations.
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quote: In North Africa, the presence of Berbers – a term to denote those populations which speak a Berber language (Camps 1980) – is well described since the Capsian (10,000–4700 years ago),
Berber languages originated in East Africa, where Afroasiatic languages were born. Therefore, the primary populations speaking proto-Berber would have been those between East Africa and the Sahara. Trying to exclusively tie the age of Berber languages to the extreme coasts of North Africa is nonsense. The languages were not "born" in the coasts of North Africa. Therefore, whatever the people of the extreme coasts of North Africa look like today, has no bearing on the origin of Berber languages and does not make populations in ancient North Africa 10,000 years ago Berber speakers. There has been no hard evidence that the populations of the extreme coasts of North Africa spoke Berber languages 10,000 years ago. All of this is pure speculation by Eurocentric clowns who want to do everything in their power to try and make Berber langues, which started in East Africa, somehow of European origin, when it is not. All of the primary expansions of Berber languages were from nomadic/semi-nomadic populations in the Sahara who ultimately originated in East Africa.
From wikipedia:
quote:Proto-Berber shows features which clearly distinguish it from all other branches of Afroasiatic, but modern Berber languages are relatively homogeneous, suggesting that whereas the split from the other known Afroasiatic branches was very ancient, on the order of 10000~9000 BP, according to grottochronological studies,[2] Proto-Berber might be as recent as 3000 BP. Louali & Philippson (2003) propose, on the basis of the lexical reconstruction of livestock-herding, a Proto-Berber 1 (PB1) stage around 7000 BP and a Proto-Berber 2 (PB2) stage as the direct ancestor of contemporary Berber languages
quote: ABSTRACT The Berber languages are relatively well-studied, and it is possible to explore their geographical extent today and in the past, and also reconstruct basic and culturalvocabulary which can be attributed to speakers of proto-Berber. However, there is a major problem reconciling this with textual and archaeological evidence. The proto-Berber we can reconstruct seems to be far to recent to match what we know from other evidence; indeed it seems to reach back to period as late as 200 AD. Textual evidence (and Canarian inscriptions) point to a period prior to 400 BC, while the most credible archaeological correlate would be the spread of pastoralism across the Sahara, pointing to the period 5-4000 BP. The paper explores this disjunction and suggests the underlying reason for it is massive language levelling in the period after 0 AD. In other words, the original speakers of Berber did indeed spread out westwards from the Nile Valley, 5-4000 years ago, but the diversity which evolved in this period was eliminated by a sociolinguistic processes which levelled divergent speech forms. Historical linguists have been wary of invoking such process until recently, but evidence is mounting for their importance in many and varied cultures, including China, Borneo and Madagascar. Hypotheses are evaluated to explain the Berber situation and it is suggested that a combination of the introduction of the camel and the establishment of the Roman limes were the key factors in creating this linguistic bottleneck.
Making it clear that trying to put the homeland of Berber languages in some coastal North African framework of Eurasian migrants is strictly nonsense.
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The information in the article pertains to the genetic make up of these particular groups as they are today and where the vast majority of them live, in the coastal regions. -not people of the region 10,000 years ago in the Saharan green period
the thread topic article does not include Alergia or Libya or Mali where there are also berber speakers The information in this article pertains to Morocco and Egypt only, the following groups
Asni Figuig Bouhriah Siwa
The last population known in the Maghreb before the berbers were the prehistoric Capsian hunter gathers beginning approximatly 12.000 years ago and and ending 8.000 years ago. No populations have been discovered in between them and the berbers although some continuity is possible. As we know a wide variety of different ethnic groups have settled in the region for the last 5-6000 years. However in terms of Y Dna E-M81 (E1b1b1b1) is a common factor spanning across all berber groups with high frequency including Tuareg with but with the exception of Egyptian Siwa who do not have high frequencies of M81
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quote:Even if archaeological and paleoanthropological records testify to the ancient (Paleolithic) human occupation of North Africa, the evolution of human groups living in that area is still unclear.
quote: M1 is a subclade of southwest Asian ancestry that moved to Africa about 40,000 to 45,000 years ago (Olivieri et al. 2006).
quote: Two other variants (489C and 10873C) also support a single origin of haplogroup M in Africa.
posted
Tajima's D and Fu's Fs, our unpublished data). An intriguing increased frequency of L0a1 in the Balanta might parallel A1-M31 and A3b2-M13 Y chromosomes in representing East African traces. Although the founder L0a1 haplotype is shared in an east-to-west corridor, the emerging lineages are exclusive of Guineans, indicating a rapid spread and local expansion after arrival.
The analysis of our data provides further evidence for the homogeneity of the Y chromosome gene pool of sub-Saharan West Africans, due to the high frequency of haplogroup E3a-M2. Its frequency and diversity in West Africa are among the highest found, suggesting an early local origin and expansion in the last 20–30 ky. Hypothesizing on the existence of an important local agricultural centre, this could have supported a demographic expansion, on an E3a-M2 background, that almost erased the pre-existing Y chromosome diversity. Its pattern of diversity within Mandenka and Balanta hints at a more marked populational growth, these people possibly related to the local diffusion of agricultural expertise. The Papel and Felupe-Djola people retain traces of their East African relatives, to which the short timescale of residence in Guinea-Bissau and higher isolation from major influences have contributed.
These minor imprints may represent movements from Sahel's more central and eastern parts, seen, for example, in the typically Ethiopian/Sudanese E3*-PN2 lineages that have reached Senegambia [2,3,5]. Y-chromosomal diversity in the population of Guinea-Bissau: a multiethnic perspective Alexandra Rosa et al.
Posts: 22234 | From: האם אינכם כילדי הכרית אלי בני ישראל | Registered: Nov 2010
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Updating Phylogeny of Mitochondrial DNA Macrohaplogroup M in India: Dispersal of Modern Human in South Asian Corridor
Adimoolam Chandrasekar et al.
quote: Abstract
To construct maternal phylogeny and prehistoric dispersals of modern human being in the Indian sub continent, a diverse subset of 641 complete mitochondrial DNA (mtDNA) genomes belonging to macrohaplogroup M was chosen from a total collection of 2,783 control-region sequences, sampled from 26 selected tribal populations of India. On the basis of complete mtDNA sequencing, we identified 12 new haplogroups - M53 to M64; redefined/ascertained and characterized haplogroups M2, M3, M4, M5, M6, M8′C′Z, M9, M10, M11, M12-G, D, M18, M30, M33, M35, M37, M38, M39, M40, M41, M43, M45 and M49, which were previously described by control and/or coding-region polymorphisms. Our results indicate that the mtDNA lineages reported in the present study (except East Asian lineages M8′C′Z, M9, M10, M11, M12-G, D ) are restricted to Indian region.The deep rooted lineages of macrohaplogroup ‘M’ suggest in-situ origin of these haplogroups in India. Most of these deep rooting lineages are represented by multiple ethnic/linguist groups of India. Hierarchical analysis of molecular variation (AMOVA) shows substantial subdivisions among the tribes of India (FST = 0.16164). The current Indian mtDNA gene pool was shaped by the initial settlers and was galvanized by minor events of gene flow from the east and west to the restricted zones. Northeast Indian mtDNA pool harbors region specific lineages, other Indian lineages and East Asian lineages. We also suggest the establishment of an East Asian gene in North East India through admixture rather than replacement.
quote:DNA polymorphisms reveal a population's genetic structure, migration and admixture in the past, susceptibility to illness and genetic causes of diseases. A phylogenetic approach is strongly recommended to avoid spurious positive associations between mtDNA mutations and diseases [1]. The pathogenic role of the mitochondrial genome requires more extensive surveys of the mtDNA sequences in different populations and patient groups. Technological improvements in DNA sequencing has made it possible to sequence complete mtDNA genome faster. Attempts have been made to reconstruct the phylogenies and prehistoric dispersal of modern humans in Europe, Africa, Oceania, East Asia, Southeast Asia and South Asia [2], [3]–[24] with complete mtDNA sequence information.
The out-of-Africa scenario [25] has hitherto provided little evidence of the precise route by which modern humans might have left Africa. Two major routes of dispersal have been hypothesized: one is through North Africa into the Levant [26], and another is through Ethiopia along South Asia [27]–[28]. The proposed northern route of initial dispersal of modem humans from Africa could not be sustained by complete and in-depth analysis of mtDNA in recent times [29]. The mitochondrial haplogroup M which was first regarded as an ancient marker of East-Asian origin [30]–[31], had been found at high frequency in India [32] and Ethiopia [33], thus raising the question of its origin. The presence of M haplogroup in Ethiopia, named M1, led to the proposal that haplogroup M originated in eastern Africa, approximately 60,000 years ago, and was carried towards Asia [34].
Contrary to the above, in 2006, Olivieri [35] reported that about 40,000 to 45,000 years ago, predominant North African clades M1 and U6 arose in southwestern Asia and moved together to Africa. Their arrival temporally overlapped the event(s) that led to the peopling of Europe by modern humans and most likely the result of the same change in the climatic conditions that allowed humans to enter in to the Levant, opening the way to the colonization of both Europe and North Africa. In the light of above, the origins of Asian M lineage in Eastern Africa became ambivalent.
Macrohaplogroup M is ubiquitous in India and covers more than 70 per cent of the Indian mtDNA lineages [28], [36]–[38]. Recent studies on complete mtDNA sequences (~187) tried to resolve the phylogeny of Indian macrohaplogroup M. As a result, M2, M3, M4, M5, M6 [28], [36], [39]–[40], M18, M25 [38], M30, [41], M31 [42], [24] M33, M34, M35, M36, M37, M38, M39, M40 [22], M41, M42 [43], M43 [23], [44], M45 [45], M48, M49, and M50 [46] haplogroups of M that was identified in India helped to a certain extent in understanding M genealogy in diversified Indian populations. In the above background, extensive sequencing of complete mtDNA of South Asia, particularly India, is essential for better understanding of the peopling of the non-African continents, and pathogenesis of diseases in various ethnic groups with different matrilineal backgrounds.
Posts: 22234 | From: האם אינכם כילדי הכרית אלי בני ישראל | Registered: Nov 2010
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The Complex and Diversified Mitochondrial Gene Pool of Berber Populations
C. Coudray1∗ , A. Olivieri2, A. Achilli2,3, M. Pala2, M. Melhaoui4, M. Cherkaoui5, F. El-Chennawi6, M. Kossmann7, A. Torroni2 and J. M. Dugoujon1
quote:Two hypotheses are discussed: one would date these common origins in the Upper Paleolithic with the ex- pansion of anatomically modern humans, from the Near East to both shores of the Mediterranean Sea; the other supports the Near Eastern origin, but would rather date it from the Neolithic, around 10,000 years ago (Ammerman & Cavalli- Sforza 1973; Barbujani et al. 1994; Myles et al. 2005; Rando et al. 1998).
quote: Common polymorphisms (i.e. those defining H and V lineages) between Berbers and south Europeans also could have been introduced or supported by genetic flows through the Straits of Gibraltar. For example, genetic exchanges could have taken place during prehistory, while European populations retreated from ice sheets and expanded from refuge, around 15,000 years ago (as evidenced by the H and U5b mitochondrial lineages).
quote:Alternatively, these exchanges could have occurred during history, with the invasion and the occupation during nearly seven centuries (from the 8th to the 15th century) of the Iberian Peninsula by Almoravide then Almohade Muslim Berber troops
Posts: 22234 | From: האם אינכם כילדי הכרית אלי בני ישראל | Registered: Nov 2010
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quote:Originally posted by Doug M: There were no "Berbers" 10,000 years ago.
quote: In North Africa, the presence of Berbers – a term to denote those populations which speak a Berber language (Camps 1980) – is well described since the Capsian (10,000–4700 years ago),
Berber languages originated in East Africa, where Afroasiatic languages were born. Therefore, the primary populations speaking proto-Berber would have been those between East Africa and the Sahara. Trying to exclusively tie the age of Berber languages to the extreme coasts of North Africa is nonsense. The languages were not "born" in the coasts of North Africa. Therefore, whatever the people of the extreme coasts of North Africa look like today, has no bearing on the origin of Berber languages and does not make populations in ancient North Africa 10,000 years ago Berber speakers. There has been no hard evidence that the populations of the extreme coasts of North Africa spoke Berber languages 10,000 years ago. All of this is pure speculation by Eurocentric clowns who want to do everything in their power to try and make Berber langues, which started in East Africa, somehow of European origin, when it is not. All of the primary expansions of Berber languages were from nomadic/semi-nomadic populations in the Sahara who ultimately originated in East Africa.
From wikipedia:
quote:Proto-Berber shows features which clearly distinguish it from all other branches of Afroasiatic, but modern Berber languages are relatively homogeneous, suggesting that whereas the split from the other known Afroasiatic branches was very ancient, on the order of 10000~9000 BP, according to grottochronological studies,[2] Proto-Berber might be as recent as 3000 BP. Louali & Philippson (2003) propose, on the basis of the lexical reconstruction of livestock-herding, a Proto-Berber 1 (PB1) stage around 7000 BP and a Proto-Berber 2 (PB2) stage as the direct ancestor of contemporary Berber languages
quote: ABSTRACT The Berber languages are relatively well-studied, and it is possible to explore their geographical extent today and in the past, and also reconstruct basic and culturalvocabulary which can be attributed to speakers of proto-Berber. However, there is a major problem reconciling this with textual and archaeological evidence. The proto-Berber we can reconstruct seems to be far to recent to match what we know from other evidence; indeed it seems to reach back to period as late as 200 AD. Textual evidence (and Canarian inscriptions) point to a period prior to 400 BC, while the most credible archaeological correlate would be the spread of pastoralism across the Sahara, pointing to the period 5-4000 BP. The paper explores this disjunction and suggests the underlying reason for it is massive language levelling in the period after 0 AD. In other words, the original speakers of Berber did indeed spread out westwards from the Nile Valley, 5-4000 years ago, but the diversity which evolved in this period was eliminated by a sociolinguistic processes which levelled divergent speech forms. Historical linguists have been wary of invoking such process until recently, but evidence is mounting for their importance in many and varied cultures, including China, Borneo and Madagascar. Hypotheses are evaluated to explain the Berber situation and it is suggested that a combination of the introduction of the camel and the establishment of the Roman limes were the key factors in creating this linguistic bottleneck.
Making it clear that trying to put the homeland of Berber languages in some coastal North African framework of Eurasian migrants is strictly nonsense.
I see,
In North Africa, the presence of Berbers – a term to denote those populations which speak a Berber language (Camps 1980) – is well described since the Capsian (10,000–4700 years ago), although this industry derived from oldest cultures.
Nevertheless, it was only during the Neolithic transition (around 6000 years ago in the Saharan areas and 5000 years ago in the Maghreb) that North Africa was incontestably marked by various cultural events.
Posts: 22234 | From: האם אינכם כילדי הכרית אלי בני ישראל | Registered: Nov 2010
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quote:Originally posted by Doug M: There were no "Berbers" 10,000 years ago.
quote: In North Africa, the presence of Berbers – a term to denote those populations which speak a Berber language (Camps 1980) – is well described since the Capsian (10,000–4700 years ago),
Berber languages originated in East Africa, where Afroasiatic languages were born. Therefore, the primary populations speaking proto-Berber would have been those between East Africa and the Sahara. Trying to exclusively tie the age of Berber languages to the extreme coasts of North Africa is nonsense. The languages were not "born" in the coasts of North Africa. Therefore, whatever the people of the extreme coasts of North Africa look like today, has no bearing on the origin of Berber languages and does not make populations in ancient North Africa 10,000 years ago Berber speakers. There has been no hard evidence that the populations of the extreme coasts of North Africa spoke Berber languages 10,000 years ago. All of this is pure speculation by Eurocentric clowns who want to do everything in their power to try and make Berber langues, which started in East Africa, somehow of European origin, when it is not. All of the primary expansions of Berber languages were from nomadic/semi-nomadic populations in the Sahara who ultimately originated in East Africa.
From wikipedia:
quote:Proto-Berber shows features which clearly distinguish it from all other branches of Afroasiatic, but modern Berber languages are relatively homogeneous, suggesting that whereas the split from the other known Afroasiatic branches was very ancient, on the order of 10000~9000 BP, according to grottochronological studies,[2] Proto-Berber might be as recent as 3000 BP. Louali & Philippson (2003) propose, on the basis of the lexical reconstruction of livestock-herding, a Proto-Berber 1 (PB1) stage around 7000 BP and a Proto-Berber 2 (PB2) stage as the direct ancestor of contemporary Berber languages
quote: ABSTRACT The Berber languages are relatively well-studied, and it is possible to explore their geographical extent today and in the past, and also reconstruct basic and culturalvocabulary which can be attributed to speakers of proto-Berber. However, there is a major problem reconciling this with textual and archaeological evidence. The proto-Berber we can reconstruct seems to be far to recent to match what we know from other evidence; indeed it seems to reach back to period as late as 200 AD. Textual evidence (and Canarian inscriptions) point to a period prior to 400 BC, while the most credible archaeological correlate would be the spread of pastoralism across the Sahara, pointing to the period 5-4000 BP. The paper explores this disjunction and suggests the underlying reason for it is massive language levelling in the period after 0 AD. In other words, the original speakers of Berber did indeed spread out westwards from the Nile Valley, 5-4000 years ago, but the diversity which evolved in this period was eliminated by a sociolinguistic processes which levelled divergent speech forms. Historical linguists have been wary of invoking such process until recently, but evidence is mounting for their importance in many and varied cultures, including China, Borneo and Madagascar. Hypotheses are evaluated to explain the Berber situation and it is suggested that a combination of the introduction of the camel and the establishment of the Roman limes were the key factors in creating this linguistic bottleneck.
Making it clear that trying to put the homeland of Berber languages in some coastal North African framework of Eurasian migrants is strictly nonsense.
I see,
In North Africa, the presence of Berbers – a term to denote those populations which speak a Berber language (Camps 1980) – is well described since the Capsian (10,000–4700 years ago), although this industry derived from oldest cultures.
Nevertheless, it was only during the Neolithic transition (around 6000 years ago in the Saharan areas and 5000 years ago in the Maghreb) that North Africa was incontestably marked by various cultural events.
That is exactly what I was referring to. How is Berber language attested to in Northern Africa 10,000 years ago? And how is it tied to the Capsian? I need to find out what Gabriel Camps actually says in this case. Sounds like they are trying to tie a tool industry in the extreme coast of North Africa from 10,000 years ago to a language which even they themselves admit didn't expand into the region until less than 5,000 years ago.
Posts: 8889 | Registered: May 2005
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The information in the article pertains to the genetic make up of these particular groups as they are today and where the vast majority of them live, in the coastal regions. -not people of the region 10,000 years ago in the Saharan green period
the thread topic article does not include Alergia or Libya or Mali where there are also berber speakers The information in this article pertains to Morocco and Egypt only, the following groups
Asni Figuig Bouhriah Siwa
The last population known in the Maghreb before the berbers were the prehistoric Capsian hunter gathers beginning approximatly 12.000 years ago and and ending 8.000 years ago. No populations have been discovered in between them and the berbers although some continuity is possible. As we know a wide variety of different ethnic groups have settled in the region for the last 5-6000 years. However in terms of Y Dna E-M81 (E1b1b1b1) is a common factor spanning across all berber groups with high frequency including Tuareg with but with the exception of Egyptian Siwa who do not have high frequencies of M81
Tukuler
multidisciplinary Black Scholar
Member # 19944
posted
quote:Originally posted by Doug M: There were no "Berbers" 10,000 years ago.
True but Berber speakers didn't materialize out of thin air and there are now no Berber speakers where it originated.
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