The term Eurasian backflow, or Eurasian back-migrations, has been used to describe several pre-Neolithic and Neolithic migration events of humans from western Eurasia back to Africa. [1]
Although preceded by many waves of earlier Homo Sapien dispersals into Eurasia, all present-day humans outside Africa descend primarily from a single expansion that occurred between 70,000–50,000 years ago. [3] [4] [5]
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. [6]
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. [7] 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. [8] 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. [7]
Outside of North Africa, about 3,000 years ago, [9] [10] or already earlier between 6,000-5,000 years ago, [11] 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. [12] 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. [9] Although Medieval events, such as the Arab expansions, also brought gene flow into various African populations, [13] these left a lower mark on North African genetic structure, relative to Neolithization. [14] [15]
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. [17] [7] 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. [18] 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] In addition to intrinsic diversity within the continent borne by population structure and isolation, influxes of Eurasian populations into Africa has been seen as a critical contributor to the existing genetic diversity. [24]
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. [25]
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." [26]
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. [27]
African Population | Time Period | Number | Proportions of Ancestry |
---|---|---|---|
Egypt, Nuerat Egyptian | 2,868 - 2,492 BC | N=1 | 90% [52] |
Tunisia, Carthaginian | 450 - 170 BCE | N=17 | [53] |
Canary Islands, Guanche | 3rd - 16th Century CE | N=40 | 89.4% [30] |
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] |
Algeria, Numidian | 1 - 700 CE | N=1 | 85.87% [56] |
Egypt, Abusir-el-Meleq Egyptian | 1388 BCE - 426 CE | N=3 | 85 - 94% [36] |
Morocco, KEB (Khef el-Baroud) | 3000 BCE | N=8 | 81.75% [57] |
Tunisia, Hergla | 5900 BP | N=1 | [58] |
Tunisia, Kerkouane Carthagian | 900 - 360 BCE | N=27 | [53] |
Tunisia, Kerkouane Carthagian | 650 - 250 BCE | N=12 | 77.33 - 97.3% [59] [60] |
Tunisia, DEK (Doukanet el Khoutifa) | 7000 - 6600 BP | N=5 | [58] |
Sudan, Kulubnarti Nubian | 650 -1000 CE | N=66 | 46 - 64% [61] |
Morocco, IAM (Ifri n'Amar) | 5000 BCE | N=7 | [57] |
Tunisia, Djebba | 8000 BP | N=2 | [58] |
Morocco, OUB (Ifri Ouberrid) | 7660 - 7506 BP | N=1 | [54] |
Algeria, ABR (Afalou Bou Rhummel) | 10,000 BP | N=1 | [58] |
Morocco, Iberomaurusian | 15,100 - 13,900 YBP | N=7 | 46 - 63.5% [43] [62] |
Sudan, Kadruka Nubian | 4033 BP | N=1 | 45.2% [63] |
Tanzania, Luxmanda | 3141 - 2890 BP | N=1 | 37.2 - 39% [64] |
Kenya, Pastoral Neolithic | 3500 - 1500 BP | N=41 | 30 - 40% [65] [8] |
Tanzania, Swahili | 1250 - 1800 AD | N=80 | 26 - 68% [66] |
Libya, Takarkori | 7,000 YA | N=2 | 20 - 30% [67] |
Combining ancient and present-day DNA data, this Neolithic demographic transformation was shown to have a greater genetic impact on present-day North Africans than later processes such as Arabization.
Previously, the West Eurasian population known to be the best proxy for this ancestry was present-day Sardinians, who resemble Neolithic Europeans genetically. However, our analysis shows that East African ancestry is significantly better modelled by Levantine early farmers than by Anatolian or early European farmers, implying that the spread of this ancestry to East Africa was not from the same group that spread Near Eastern ancestry into Europe.
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: CS1 maint: numeric names: authors list (link)The Nubian, Arab, and Beja populations of northeastern Africa roughly display equal admixture fractions from a local northeastern African gene pool (similar to the Nilotic component) and an incoming Eurasian migrant component (53) (Figure 3).
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: CS1 maint: numeric names: authors list (link)At Kerkouane, a Carthaginian town on the Cap Bon peninsula in Tunisia (see extended description in Materials), we observe a highly heterogeneous population, spanning across the PCA space in Fig. 3 from modern Mozabite populations to modern Sicilian populations, consisting of three primary genetic clusters.
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