Eurasian backflow

Pre-Neolithic and Neolithic human migration events From Wikipedia, the free encyclopedia

"Eurasian backflow" or "Eurasian back-migrations" refers to several waves of early modern-human migrations before and during the Neolithic, when some groups of anatomically modern humans (contemporaneous Homo sapiens) returned to Africa from parts of Eurasia.[1] Modern humans had begun dispersing into Eurasia from Africa during the Middle Paleolithic. Today, all humans outside of Africa descend primarily from a single expansion that occurred 70,000–50,000 years ago.[2][3][4]

Modern humans who dispersed into Eurasia came into contact with some groups of archaic humans, such as Neanderthals, and eventually interbred with them. As a result of this phenomenon, which continued until the Upper Paleolithic, those who returned from Eurasia played a significant role in introducing Neanderthal DNA to the African modern human genome, particularly among North Africans. Thus, in addition to intrinsic diversity within the continent borne by population structure and isolation, these historical influxes of Eurasian populations into Africa are regarded as a critical contributor to the existing genetic diversity.[5]

Migration history

Pre-Neolithic and Neolithic migration events in Africa.[6]

Beginning around 30,000-15,000 years ago, a population closely related to Paleolithic individuals from Dzudzuana Cave in Georgia expanded from the Near East back into Northern Africa, spreading in connection with microlithic-backed bladelet technologies that would later become ubiquitous across North Africa. This diffusion contributed up to 55% of the genetic ancestry of the Iberomaurusian culture, and introduced the mitochondrial haplogroups M1 and U6 to the region, which along with the aforementioned genetic ancestry, define the "autochthonous/typical North African" or "Maghrebi" component, as it is referred to in genetic studies.[7]

Approximately 8,000 years ago, populations of the Neolithic Levant expanded into North Africa via the Sinai Peninsula. This diffusion was rooted in wider displacements of people in the wake of the 6200 BC climatic event, and coincided with the introduction of Southwest Asian domesticates such as sheep, goats, and cattle into Africa, leading to the rise of Saharan cattle pastoralism and the appearance of Ashakar Ware pottery in the Maghreb.[8] This expanding population interacted and admixed with local ancestries, as seen in both Skhirat-Rouazi individuals of Morocco and eastern African pastoralist groups of the Pastoral Neolithic.[9] This diffusion may have also provided the demographic and cultural conditions for the early dispersal of Afro-Asiatic languages across North Africa, whose known branches of Cushitic, Chadic, Egyptian, and Berber correspond strikingly with the geography of these Neolithic expansions.[8]

Outside of North Africa, about 3,000 years ago,[10][11] or already earlier between 6,000-5,000 years ago,[12] farmers from Anatolia and the Near East migrated into other regions. Signs of this migration can be found in the genomes of contemporary peoples from all over East Africa. Moreover, analysis has also recognized that some of the Eurasian ancestry in Northeast Africa could possibly pre-date agriculture, from around ~12-23 ka.[13] Significant Eurasian ancestry is found among populations of Eastern Africa, and among specific ethnic groups of the Horn of Africa, Northern Sudan, the Sahel region, as well as among the Malagasy people of Madagascar. West Eurasian ancestry thus arrived in Africa first during the Palaeolithic, then followed by several later Neolithic, Chalcolithic, and Bronze Age migration events.[10] Although Medieval events, such as the Arab expansions, also brought gene flow into various African populations,[14] these left a lower mark on North African genetic structure, relative to Neolithization.[15][16]

Map of major prefarming population stratification across the African continent.[17]

The Neolithic populations migrating back-to-Africa were said to be similar to the Neolithic farmers who had brought agriculture from Anatolia to Europe about 7,000 years ago, and whose demographic legacy is most closely preserved in present-day Sardinians, their contemporary descendants.[1] Although, other studies have revealed some genetic distinctions between the populations that brought farming into Europe and the Maghreb, and the Levantine related groups that spread southward into East Africa.[18][8] A study from 2020 inferred two sources for the spread of Eurasian admixture in Northeastern Africa, with one being associated with pastoralism, involving groups originating from the Levant and North Africa.[6] Further research has shown that the back-migration into the region was a complex process, identifying multiple origins for the Eurasian component in Northeast African groups today.[19][20]

A report in November 2015 on a 4,500-year-old Ethiopian genome[21][22] had originally overestimated the genetic influence of the Eurasian backflow, claiming that signs of the migration could be found in genomes all over Africa. This mistaken claim was based on a data-processing error and was corrected in February 2016. The West Asian admixture was only predominant in the populations of the Horn of Africa, in particular Ethiopian highlanders, and less relevant or absent in the genetic makeup of West and Central Africans.[23]

Neanderthal admixture and Eurasian ancestry

An investigation in 2012 discovered that unlike most sub-Saharan Africans, North Africans have similar levels of Neanderthal DNA to South Europeans and West Asians, which is pre-Neolithic in origin, rather than via any recent admixture, as the Neanderthal's genetic signals were higher in populations with an autochthonous 'back-to-Africa' genomic component that arrived 12,000 years ago. These Neanderthal genomic traces do not mark a division between Africans and non-Africans, but rather a division between sub-Saharan Africans and the rest of the modern human groups, including those from North Africa.[24]

In 2016, researchers recognized that the Neanderthal ancestry in African populations, strongly corresponds with the levels of Western Eurasian ancestry. The geneticists elaborated that: "Neanderthal ancestry is not expected in Africa, yet today many Africans carry Neanderthal-derived alleles. The plot shows that the Neanderthal ancestry proportion in Africans is correlated with gene flow from Eurasians. For example, knowing that today Eurasians carry ~2% of Neanderthal ancestry, we observed that East Africans (Ethiopians) had ~1% Neanderthal ancestry and ~50% Eurasian ancestry. Correspondingly, Near Easterners showed a decline in Neanderthal ancestry proportional to their levels of African ancestry."[25]

Human dispersals within Africa

Chen, Lu's publication found back-migrations contributed to the signal of Neanderthal ancestry in Africans. Data indicated that back-migrations giving Neanderthal sequences came after the split of Europeans and East Asians, from populations related to the European lineage. The overlap of this ancestral European ancestry and Neanderthal segments was highly significant.[26]

Average Eurasian frequencies in African populations

More information African Population Group, Linguistic Affiliation ...
Estimates in Modern Africans
African Population Group Linguistic Affiliation Proportions of Ancestry
Afrikaner Germanic 97.9%[27]
Rashaida Arab Semitic 97.92%[28]
Canarian Romance 97.4%[29]
Tunisian Jew Semitic 96.49%[30]
Moroccan Jew Semitic 96.05%[30]
Libyan Jew Semitic 95.76%[30]
Chaoui Berber 85.12 - 87.51%[31]
Moroccan Semitic 80%[32]
Tunisian Semitic 79.37 - 81%[30][33]
Egyptian Semitic 79 - 86%[32][34][35]
Figuig Berber Berber 78.98%[28]
Bouhria Berber Berber 78.68%[28]
Mozabite Berber 77.33 - 82.39%[30][32][31]
Asni Berber Berber 76.99%[28]
Algerian Semitic 76.86 - 82.45%[30][31]
Libyan Semitic 76%[36]
Saharawi Semitic 74.32 - 84.9%[30][36]
Tigrayan Ethio-Semitic 50 - 50.4%[37][10]
Beja Cushitic 50%[38]
Amhara Ethio-Semitic 49.2 - 54%[10][34][39]
Ethiopian Jew Semitic 46.4 - 55.12%[40][30]
Afar Cushitic 46.0%[10]
Kababish Arab Semitic 45.99%[28]
Oromo Cushitic 41.6 - 51%[10][39]
Agaw Cushitic 41 - 50%[41]
Sudanese Arab Semitic 40 - 52.73%[14][28]
Nubian Nubian 39.41 - 54.74%[14][28]
Somali Cushitic 38.4 - 44%[10][42][30][39]
Cape Verdean Romance 36 - 54%[43][44]
Wolayta Damotic 34.1 - 43%[10][39]
Samburu Nilotic 32.19%[28]
Datooga Nilotic 32.17%[30]
Malagasy Austronesian 30 - 68%[45][46]
Baggara Arab Semitic 25.57 - 33.6%[28]
Fulani Atlantic 21.4 - 29%[47][48]
Toubou Saharan 20 - 31.4%[25][36]
Maasai Nilotic 18.9 - 26.06%[10][30]
Turkana Nilotic 18.33%[28]
Kikuyu Bantu 16.4 - 18.8%[40][30]
Aari Aroid 15.7 - 18.2%[10]
Sandawe Sandawe 15.4%[10]
Nama Khoi 14%[10]
Zaghawa Saharan 12.58%[28]
Kalenjin Nilotic 10.99 - 12.86%[40][28]
Kanembu Saharan 10.97%[28]
Hadza Hadza 6.4 - 12.71%[10][30]
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More information African Population, Time Period ...
Estimates in Ancient African Samples
African Population Time Period Number Proportions of Ancestry
Egypt, Abusir-el-Meleq Egyptian 1388 BCE - 426 CE N=3 94 - 96%[35][49][50]
Tunisia, Carthaginian 450 - 170 BCE N=17 92.62 - 100%[51]
Tunisia, Kerkouane Carthaginian 900 - 250 BCE N=27 92.62 - 100%[51][52]
Egypt, Nuerat Egyptian 2,868 - 2,492 BC N=1 91.86%[49][50]
Algeria, Numidian 1 - 700 CE N=3 89.4 - 100%[53]
Canary Islands, Guanche 3rd - 16th Century CE N=40 89.4%[29]
Morocco, SKH (Skhirat-Rouazi) 6733 - 6121 BP N=3 87%[54][55]
Morocco, KTG (Kaf Taht el-Ghar) 7429 - 6945 BP N=4 86%[54][55]
Morocco, KEB (Khef el-Baroud) 3000 BCE N=8 81.75%[56]
Tunisia, Hergla 5900 BP N=1 ?[57]
Tunisia, DEK (Doukanet el Khoutifa) 7000 - 6600 BP N=5 ?[57]
Tunisia, Djebba 8000 BP N=2 ?[57]
Morocco, IAM (Ifri n'Amar) 5000 BCE N=7 ?[56]
Morocco, Iberomaurusian 15,100 - 13,900 YBP N=7 60 - 70%[58][59][60]
Algeria, ABR (Afalou Bou Rhummel) 10,000 BP N=1 ?[57]
Morocco, OUB (Ifri Ouberrid) 7660 - 7506 BP N=1 ?[54]
Sudan, Kulubnarti Nubian 650 -1000 CE N=66 46 - 64%[61]
Sudan, Kadruka Nubian 4033 BP N=1 45.2%[62]
Tanzania, Luxmanda 3141 - 2890 BP N=1 37.2 - 39%[63]
Kenya, Pastoral Neolithic 3500 - 1500 BP N=41 30 - 40%[64][10]
Tanzania, Swahili 1250 - 1800 AD N=80 26 - 68%[65]
Libya, Takarkori 7,000 YA N=2 25%[58]
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