Upper Paleolithic ties between E. Africa & Europe

[Noah]

Thought I'd share the following interesting passage on the morphological similarities between the Upper Paleolithic peoples of East Africa and Europe, drawn from the anthropologist Sonia Cole's classic The Prehistory of East Africa.

Comparison between Upper Palaeolithic man in Europe and Africa

"Before leaving Upper Palaeolithic man in East Africa, we should glance briefly at his contemporaries in Europe living during the last glacial period. The Upper Palaeolithic people who invaded Europe after the first phase of the Wurm glaciation arrived in a series of waves, probably from the direction of Palestine. First came the Combe Capelle type, makers of the Chatelperronian culture (which is similar to the Upper Kenya Capsian and to the later Capsian of North Africa). These are the earliest known representatives of Homo sapiens in Europe, apart from the Swanscombe and Fontechevade fossils. They were followed by the Aurignacians of Grimaldi and then by the robust Cro-Magnons, associated with Aurignacian and Gravettian industries. In the Cro-Magnons, Coon sees an admixture of Neanderthal blood, possibly springing from crosses as may have occurred at Mount Carmel. The Cro-Magnons are divisible into a dolichocephalic eastern or Predmost type (which is very close to Combe Capelle), and a broad-headed western type, the Cro-Magnons proper. After them came the Magdalenians known from Chancelade. All these different types may have arrived in Europe within a few thousand years.

The skulls from Gamble's Cave, the Naivasha rock shelter, and Olduvai are very similar to the large-brained Combe Capelle type. Cro-Magnon man is less modern in appearance than the earlier Combe Capelle people; he resembles the 'Mechta' type of North Africa. The conclusion is that Upper Palaeolithic peoples in Africa differed hardly at all in appearance (or in the form of implements that some of them made), from their counterparts in Europe."

Cole states elsewhere that the Negroid element was only later introduced to East Africa with the expansion of Bantu and Nilotic tribes into the region, as does W.W. Howells and most other scholars. On this she writes:

"One of the most striking facts to emerge is the comparatively late appearance of the Negroid type -- in the Mesolithic Khartoum, but not before Neolithic times in Kenya. Apart from the proto-Bushman skull from Singa, and the specialized proto-Australoid from Eyasi, the Upper Paleolithic people known from East Africa were of the Caucasoid stock, or proto-Hamites."

The lack of morphological ties between Upper Paleolithic Europeans and modern Sub-Saharan Africans was also later confirmed by Loring Brace. Only populations with predominant Caucasoid affinities, such as Hamitic peoples (Somalis), Nubians, South Asians and modern Europeans, in varying degrees share links with these ancient specimens (more on that here):

What's especially interesting from a contemporary perspective about Cole's material is that the Eurasian skeletal affinities and similarities in lithic/stone industries that she and others identify as linking the prehistoric peoples in East Africa and Europe during roughly the same period of time have now also been confirmed by genetic data.

That is, modern European maternal DNA/mtDNA is usually divided into two groups: one associated with Europe's older Upper Paleolithic hunter-gatherer populations, and the other associated with more recent Neolithic migrant farmers from the Near East. Maternally, most modern Europeans tend to descend from Upper Paleolithic hunter-gatherers (paternally, most are descended from the Near Eastern agriculturalists).

"The ancestry of modern Europeans is a subject of debate among geneticists, archaeologists, and anthropologists. A crucial question is the extent to which Europeans are descended from the first European farmers in the Neolithic Age 7500 years ago or from Paleolithic hunter-gatherers who were present in Europe since 40,000 years ago. Here we present an analysis of ancient DNA from early European farmers. We successfully extracted and sequenced intact stretches of maternally inherited mitochondrial DNA (mtDNA) from 24 out of 57 Neolithic skeletons from various locations in Germany, Austria, and Hungary. We found that 25% of the Neolithic farmers had one characteristic mtDNA type and that this type formerly was widespread among Neolithic farmers in Central Europe. Europeans today have a 150-times lower frequency (0.2%) of this mtDNA type, revealing that these first Neolithic farmers did not have a strong genetic influence on modern European female lineages. Our finding lends weight to a proposed Paleolithic ancestry for modern Europeans."

www2.webmatic.it/workO/s/113/pr-926-file_it-Haak%20aDNA%20Science.pdf

Both general maternal/mtDNA types (Upper Paleolithic & Neolithic) are present in the Hamitic populations of North and Northeast Africa. This is consistent with the Hamitic peoples' proposed descent from older Paleo-Hamitic as well as later Proto-Hamitic waves of expansion of West Eurasian populations from the Middle East.

Neolithic Eurasian mtDNA clades

The Eurasian clades that were introduced into Europe in the Neolithic by Near Eastern agriculturalists (such as haplogroup N1) are also relatively common in descendants of the Later Proto-Hamitic pastoralists, such as the Hamitic groups in the Horn and East Africa. Amongst the Cushitic-speaking Rendille herders (who are also partly of Paleo-Hamitic descent; see below), haplogroup N1a represents almost 10% of their maternal lineages.

Despite being rare in modern populations, N1a was quite common in parts of Neolithic Europe. Subsequent migrations appear to have diluted its presence in modern European populations. Multiple recent studies on prehistoric European skeletons, however, have found that many specimens from the period belonged to the clade (around 25% of the fossils in the paper below).

"The most striking result is that 6 of the 24 Neolithic skeletons are of the distinctive and rare N1a branch. For verification, we sequenced 517 clones derived from independent extractions from different parts of the six individuals. All six showed the suite of mutations characteristic of the N1a lineage[...] The high frequency of our Neolithic N1a lineages is not a local phenomenon but is widespread in the LBK area: Independently sampled locations in Hungary and Germany, over 800 km apart, each yielded one or more N1a types (Fig. 1)[...]

The results from the Neolithic sample show that other mtDNA lineages considerably diluted the mtDNA pool of these early Neolithic populations, so that the frequency of N1a in modern Europeans is 150 times lower than in our sample of the first Central European farmers. This is incompatible with the idea that modern Central Europeans—and by implication other Europeans beyond the LBK/AVK area—derive their maternal lineages purely from the earliest farmers of that region. Within the current debate on whether Europeans are genetically of Palaeolithic or Neolithic origin, and leaving aside the possibility of significant post-Neolithic migration, our data lend weight to the arguments for a Palaeolithic origin of Europeans."

www2.webmatic.it/workO/s/113/pr-926-file_it-Haak%20aDNA%20Science.pdf

To this end, note in the plot below the continued biological affinities between the Hamitic peoples of North and Northeast Africa and many populations of Neolithic Europe:

Upper Paleolithic Eurasian mtDNA clades

Upper Paleolithic Eurasian mtDNA clades such as haplogroup I are especially common in Cushitic-speaking groups like the El Molo i.e. groups that are mostly associated with the older Paleo-Hamites responsible for the pre-Bantu expansion Upper Paleolithic Kenya Capsian culture. Although haplogroup I is now infrequent in Europe, it was apparently much more common in ancient Scandinavia before later migrations into the region significantly reduced its presence.

"The overall occurrence of haplogroups did not deviate from extant Scandinavians, however, haplogroup I was significantly more frequent among the ancient Danes (average 13%) than among extant Danes and Scandinavians (~2.5%) as well as among other ancient population samples reported. Haplogroup I could therefore have been an ancient Southern Scandinavian type “diluted” by later immigration events."

It's amazing how accurate a lot of the old anthropology has proven itself to be despite the fact that scholars back then often had a much more modest set of tools and data to work with.

[Noah]

More evidence published this past year of the widespread distribution in Neolithic Europe of the Near Eastern mtDNA lineage N1a, a clade that today is most common in the Middle East and amongst Cushitic-speaking Hamitic groups in East Africa:

"To extend existing knowledge of the mitochondrial European Neolithic gene pool, we examined six samples of human skeletal material from a French megalithic long mound (c.4200 cal BC). We retrieved HVR-I sequences from three individuals and demonstrated that in the Neolithic period the mtDNA haplogroup N1a, previously only known in central Europe, was as widely distributed as western France."

onlinelibrary.wiley.com/doi/10.1002/ajpa.21376/abstract

Yemen also appears to have among the highest haplotype diversities of N1a, though greater frequencies of the clade have since been found in Central Europe and among Hamitic groups in East Africa.

"Yemeni N1a sequences[...] display a high level of haplotype (h=0.89) and nucleotide (ρ=2.75±1) diversity, combined with the highest frequency (6.9%) of this haplogroup reported so far."

www.ncbi.nlm.nih.gov/pmc/articles/PMC1182106/

[Noah]

Still more evidence published this summer of the widespread distribution in Neolithic Europe of the Near Eastern mtDNA lineage N1a, a clade that today is most often associated with West Eurasian-descended Hamitic people in North & Northeast Africa:

"It has been established, based on ancient DNA analysis of early Neolithic skeletons from various Central - European sites, that the frequency of N1a mtDNA haplogroup was 150 times more prevalent among first Neolithic farmers than in contemporary European populations (only 0.2 %), suggesting that first Neolithic farmers did not have strong impact on maternal genetic heritage of modern Europeans. This Neolithic type of N1a haplogroup belong to European sub - branch defined by control region 16147A variant. Here we present the influence of first Neolithic farmers in Southern – Eastern Europe due to finding of N1a mtDNA haplogroup in populations of Croatia and Adriatic Islands as well as Herzegovina, and complete lack of this haplogroup among populations of Serbia, Macedonia, Bosnia, Kosovo and Montenegro. We also present very interesting finding of even more ancestral lineage of N1a haplogroup in Croatian mainland and on two different Islands (Pag and Cres) belonging to African/South Asian branch characterized by control region motif 16147G. This branch is more prevalent in Arabian Peninsula and northern Africa and limitedly present also around Israel, Iran, Turkey and Greece and it is very rare in any European populations. On the Croatian island of Cres, this very unusual N1a lineage of the unknown origin has been recorded. Due to the influence of genetic drift the prevalence of this lineage is around 9.24% in total Island population. To the best of our knowledge this is the most prevalent finding of this haplogroup in any investigated population so far."

www.isabs.hr/PDF/ISABS_2011_abstract_book_web.pdf

[Noah]

Yet more evidence was published earlier this year of the widespread distribution in Neolithic Europe of the Eurasian mtDNA lineage N. In this case, the basal paragroup N*, which is ancestral to the N1 clade that's especially common amongst Hamitic people in North Africa and the Horn. The maternal haplogroup X1, whose parent clade X descends from haplogroup N, was also observed at considerable frequencies. Like haplogroup N1 discussed above, X1 is today quite rare and largely restricted to North Africa, the Horn and the Near East. The ancient DNA was extracted from archaeological sites in Iberia associated with the Cardial/Cardium Pottery/Impressed Ware culture, which is the main culture of the Mediterranean Neolithic, extending from the Adriatic to the Maghreb. The authors attribute the presence of the lineages to population movements from the Near East, the same general area as where the various waves of Hamitic migrants also expanded into Africa from:

"The Neolithic transition has been widely debated particularly regarding the extent to which this revolution implied a demographic expansion from the Near East. We attempted to shed some light on this process in northeastern Iberia by combining ancient DNA (aDNA) data from Early Neolithic settlers and published DNA data from Middle Neolithic and modern samples from the same region. We successfully extracted and amplified mitochondrial DNA from 13 human specimens, found at three archaeological sites dated back to the Cardial culture in the Early Neolithic (Can Sadurní and Chaves) and to the Late Early Neolithic (Sant Pau del Camp). We found that haplogroups with a low frequency in modern populations—N* and X1—are found at higher frequencies in our Early Neolithic population (∼31%). Genetic differentiation between Early and Middle Neolithic populations was significant (FST∼0.13, P less than 10−5), suggesting that genetic drift played an important role at this time. To improve our understanding of the Neolithic demographic processes, we used a Bayesian coalescence-based simulation approach to identify the most likely of three demographic scenarios that might explain the genetic data. The three scenarios were chosen to reflect archaeological knowledge and previous genetic studies using similar inferential approaches. We found that models that ignore population structure, as previously used in aDNA studies, are unlikely to explain the data. Our results are compatible with a pioneer colonization of northeastern Iberia at the Early Neolithic characterized by the arrival of small genetically distinctive groups, showing cultural and genetic connections with the Near East."

onlinelibrary.wiley.com/doi/10.1111/j.1365-294X.2011.05361.x/abstract

[Noah]

Still more evidence has just been published on the repeated penetration of West Eurasians into Africa. Based on mitochondrial analysis, Fernandes et al. (2012) indicate a succession of migrations from Arabia to the Horn region through the Bab el-Mandeb strait. These movements brought the aforediscussed West Eurasian mtDNA haplogroups N1 and I, clades which are today most common in the Cushitic-speaking populations of East Africa. Since these migrations involved maternal lineages (which are transmitted by females only), we are looking at new settlements of entire populations, women and men alike: the women that passed on those mtDNA clades and their male companions/protectors who accompanied them.

"The presence of archaeological sites in the Gulf basin demonstrates a long tradition of human occupation.9 However, neither direct cultural influences from the Levant nor any African influence has been detected in the Upper Palaeolithic (Late Pleistocene) lithics observed in eastern Arabia, pointing to a local development of cultural techniques.9,47 Curiously, however, the fact that some of the branches studied here include deep lineages in eastern Africa (haplogroups I, N1a, and N1f) shows that migration back to Africa occurred a number of times between 15 and 40 ka ago.

The hypothesized Gulf Oasis9 appears to be the most likely locus of the earliest branching of haplogroup N, including the three relict basal N(xR) haplogroups studied here, as well as the major Eurasian haplogroup R. Time estimates, frequencies, and genetic diversities reported here for these haplogroups are often similar between the Levant and Arabia, challenging the hypothesis of longterm isolation between these two regions. The other two refugia identified in the south and southwest of the Peninsula might have acted as a corridor for migrations west, back toward eastern Africa."

[egypt1101]

The findings of haplogroup N (above study) seem to correlate with this 2004 study, they mention Aksum which I found interesting...


“Nine distinct subclades, including three newly defined ones, were found to characterize entirely the variation of Ethiopian and Yemeni L3 lineages. Both Ethiopians and Yemenis contain an almost-equal proportion of Eurasian-specific M and N and African-specific lineages and therefore cluster together in a multidimensional scaling plot between Near Eastern and sub-Saharan African populations… Lineages that belong to haplogroup N that cover virtually all mtDNA sequences in western Eurasia (Richards et al. 2000) show substantial frequencies both in the Yemeni (44%) and Ethiopian (31%) mtDNA pools. In this respect, Ethiopians differ explicitly from most other sub-Saharan African populations studied thus far. Within Ethiopia, the frequency of N lineages is significantly higher (P<.05) in samples that originate from its northern territory (48%), which was the center of the Aksum kingdom, than among other Ethiopians, mostly originating from the south-central part of the country (27%). At the same time, there was no significant difference in the proportions of haplogroup N between the Semitic and Cushitic linguistic groups in our sample—for example, between Amharas and Oromos… Their elevated frequency and uniform presence among major language groups in Ethiopia (table 1) suggests that these derived lineages may represent a relatively old introgression of lineages to the Ethiopian mtDNA pool from the Near East…Haplogroup (preHV)1 is by far the most frequent (10.4%) subclade in the Ethiopian N cluster (fig. 2B). The majority of the Ethiopian (preHV)1 lineages match or derive from founder haplotypes common to Near Eastern, southern Caucasian, and North African populations (Krings et al. 1999; Metspalu et al. 1999; Richards et al. 2000; Kivisild et al. 2003b)… Three of the five haplogroup J lineages in Ethiopians share a distinct HVS-I motif, 16069-16126-16193-16300-16309 (J1c), that is characteristic of J sequences in populations from the southern Caucasus, the Near East, and North Africa (Di Rienzo and Wilson 1991; Richards et al. 2000; Brakez et al. 2001; Maca-Meyer et al. 2001; Plaza et al. 2003)… All Ethiopian and Yemeni haplogroup T sequences clustered with either T1 or T2 subclades, consistent with the classification of all existing European T coding-region sequences (Ingman et al. 2000; McMahon et al. 2000; Finnilä et al. 2001; Herrnstadt et al. 2002; Coble et al. 2004)… All eight Ethiopian U6 samples descend from the major U6a1 founder (fig. 2B), which is spread from the Near East to northwestern Africa at appreciable frequencies (Maca-Meyer et al. 2003). Their absence in Yemen suggests that these U6 lineages have likely penetrated to Ethiopia from the north rather than by the sea route from Arabia. (“Ethiopian Mitochondrial DNA Heritage: Tracking Gene Flow Across and Around the Gate of Tears” Toomas Kivisild et al., 2004)

The Aksum kingdom dna might be worth investigating, I haven't looked recently, but when I searched a couple years ago, didn't find anything.

[Noah]

Haplogroup N is indeed also found in the Semitic-speaking populations of Ethiopia, not just the Cushitic speaking ones. In Kivisild et al. (2004), the Semitic-speaking Tigrai population in northern Ethiopia actually had a higher percentage of West Eurasian mtDNA clade carriers than even the Yemeni sample set.

However, the presence of Eurasian maternal clades in Ethiopia and elsewhere in East Africa is in fact not primarily associated with the Axumite kingdom or recent migration. This is due to two main reasons.

First, certain West Eurasian mtDNA clades, such as haplogroup I, are most common in Cushitic-speaking populations like the El Molo (they actually have a world high of the haplogroup at ~22%). These are groups that are mostly associated with the older Paleo-Hamites responsible for the pre-Bantu expansion, Upper Paleolithic Kenya Capsian culture. They're quite mixed now since they live in a Negroid majority area (Kenya), but the West Eurasian maternal lineages that many of their constituents still carry reveal their origins quite well.

Second, Ethiopia's Cushitic-speaking communities (who are linked with other kingdoms like Zagwe) actually on average possess a slightly higher number of Eurasian clade bearers than the country's Semitic-speaking populations (who are mainly associated with Axum). For instance, in Scott et al. (2005)'s Figure 4 below, about 50% of the Cushitic-speaking samples are Eurasian clade-bearers versus about 45% of the Semitic speakers (the clades Scott et al. provisionally labeled "E1" and "E2" are Eurasian in affinity, with E2 likely representing haplogroup N derivatives):

This is why, using SAMOVA genetic analysis, Stefflova et al. clustered their Horner mtDNA samples with those in Egypt i.e. because maternally the populations are not that much different.

"Africa was divided based on geography combined in a few cases with ethnicity (in the case of Pygmy hunter gatherers and Bantu speakers). SAMOVA was used to eliminate the following outlier groups: West Pygmy (Cameroon, C.A.R., and Gabon), Khoisan speakers (South Africa and Botswana), East Pygmy group Mbuti (D.R.C.), and Moroccan sample (mainly Berbers from North Africa). After removing outliers, the remaining states were divided using SAMOVA into 4 groups: WC/SW Bantu speakers from Angola, Cameroon, Gabon and Equatorial Guinea, E/SE sample from Kenya and Mozambique Bantu speakers, NE/E sample from Egypt, Sudan, Eritrea, Ethiopia, and Somalia, and W/WC countries after excluding Bantu speakers and Pygmy hunter-gatherers[...] MDS plot showing the mtDNA variation-based genetic distances between African populations after the outliers[...] were excluded from the calculations. This plot shows the general structure of the remaining regions (with highlighted W/WC clustering) and their relationship to the admixed African American populations, depicted using only the African portion of their ancestry."

www.plosone.org/article/info%3Adoi%2F10.1371%2Fjournal.pone.0014495

Those are some of the many reasons why one can't speak of those West Eurasian maternal clades as being "intrusive" into said Horn populations anymore than one can say the same for the Egyptian or South Arabian communities. What we are looking at are entire West Eurasian populations -- women and their male companions/protectors -- that repeatedly entered Northeast Africa, formed new settlements, and whose actual descendants today have retained much of their ancestors' language, culture, somatic characteristics and genetics (Y DNA, mtDNA and autosomal DNA) despite later periods of Sub-Saharan admixture.