Genetic history of Eastern Africa

The genetic history of Eastern Africa encompasses the genetic history of the people of Eastern Africa. The Sahara served as a trans-regional passageway and place of dwelling for people in Africa during various humid phases[1][2][3] and periods throughout the history of Africa.[4][5]

Archaic Human DNA

Further information: Interbreeding between archaic and modern humans § Archaic African hominins

While Denisovan and Neanderthal ancestry in non-Africans outside of Africa are more certain, archaic human ancestry in Africans is less certain and is too early to be established with certainty.[6]

Ancient DNA

Ethiopia

At Mota, in Ethiopia, an individual, estimated to date to the 5th millennium BP, carried haplogroups E1b1 and L3x2a.[7][8] The individual of Mota is genetically related to groups residing near the region of Mota, and in particular, are considerably genetically related to the Aari people, especially the blacksmith caste of that group.[9][10]

Kenya

At Jawuoyo Rockshelter, in Kisumu County, Kenya, a forager of the Later Stone Age carried haplogroups E1b1b1a1b2/E-V22 and L4b2a2c.[11][12]

At Ol Kalou, in Nyandarua County, Kenya, a pastoralist of the Pastoral Neolithic carried haplogroups E1b1b1b2b2a1/E-M293 and L3d1d.[11][12]

At Kokurmatakore, in Marsabit County, Kenya, a pastoralist of the Pastoral Iron Age carried haplogroups E1b1b1/E-M35 and L3a2a.[11][12]

At White Rock Point, in Homa Bay County, Kenya, there were two foragers of the Later Stone Age; one carried haplogroups BT (xCT), likely B, and L2a4, and another probably carried haplogroup L0a2.[11][12]

At Nyarindi Rockshelter, in Kenya, there were two individuals, dated to the Later Stone Age (3500 BP); one carried haplogroup L4b2a and another carried haplogroup E (E-M96, E-P162).[13][14]

At Lukenya Hill, in Kenya, there were two individuals, dated to the Pastoral Neolithic (3500 BP); one carried haplogroups E1b1b1b2b (E-M293, E-CTS10880) and L4b2a2b, and another carried haplogroup L0f1.[13][14]

At Hyrax Hill, in Kenya, an individual, dated to the Pastoral Neolithic (2300 BP), carried haplogroups E1b1b1b2b (E-M293, E-M293) and L5a1b.[13][14]

At Molo Cave, in Kenya, there were two individuals, dated to the Pastoral Neolithic (1500 BP); while one had haplogroups that went undetermined, another carried haplogroups E1b1b1b2b (E-M293, E-M293) and L3h1a2a1.[13][14]

At Kakapel, in Kenya, there were three individuals, one dated to the Later Stone Age (3900 BP) and two dated to the Later Iron Age (300 BP, 900 BP); one carried haplogroups CT (CT-M168, CT-M5695) and L3i1, another carried haplogroup L2a1f, and the last carried haplogroup L2a5.[13][14]

At Panga ya Saidi, in Kenya, an individual, estimated to date between 496 BP and 322 BP, carried haplogroups E1b1b1b2 and L4b2a2.[15]

Kilifi

At Kilifi, Mtwapa, in Kenya, an individual, dated between 1250 CE and 1650 CE, carried haplogroup L3b1a1a.[16]

At Kilifi, Mtwapa, in Kenya, an individual, dated between 1250 CE and 1650 CE, carried haplogroup L0a1b2a.[16]

At Kilifi, Mtwapa, in Kenya, an individual, dated between 1250 CE and 1650 CE, carried haplogroups J1a2a1a2d2b~ and L3e3a.[16]

At Kilifi, Mtwapa, in Kenya, an individual, dated between 1250 CE and 1650 CE, carried haplogroup L3b1a11.[16]

At Kilifi, Mtwapa, in Kenya, an individual, dated between 1250 CE and 1650 CE, carried haplogroup L3b1a1.[16]

At Kilifi, Mtwapa, in Kenya, an individual, dated between 1250 CE and 1650 CE, carried haplogroups J1a2a1a2d2b~ and L0a2a1a2.[16]

At Kilifi, Mtwapa, in Kenya, an individual, dated between 1250 CE and 1650 CE, carried haplogroups J1a2a1a2d2b~ and L2a1a.[16]

At Kilifi, Mtwapa, in Kenya, an individual, dated between 1250 CE and 1650 CE, carried haplogroups J1a2a1a2 and L3f1b1a1.[16]

At Kilifi, Mtwapa, in Kenya, an individual, dated between 1250 CE and 1650 CE, carried haplogroup L3b1a1a.[16]

At Kilifi, Mtwapa, in Kenya, an individual, dated between 1250 CE and 1650 CE, carried haplogroup L0a2.[16]

At Kilifi, Mtwapa, in Kenya, an individual, dated between 1250 CE and 1650 CE, carried haplogroup L3f1b4a1.[16]

At Kilifi, Mtwapa, in Kenya, an individual, dated between 1250 CE and 1650 CE, carried haplogroup L3e3.[16]

At Kilifi, Mtwapa, in Kenya, an individual, dated between 1250 CE and 1650 CE, carried haplogroup L0a2a2a1.[16]

At Kilifi, Mtwapa, in Kenya, an individual, dated between 1250 CE and 1650 CE, carried haplogroups J and L3b1a1.[16]

At Kilifi, Mtwapa, in Kenya, an individual, dated between 1250 CE and 1650 CE, carried haplogroup L1b1a.[16]

At Kilifi, Mtwapa, in Kenya, an individual, dated between 1250 CE and 1650 CE, carried haplogroup L2a1f1.[16]

At Kilifi, Mtwapa, in Kenya, an individual, dated between 1250 CE and 1650 CE, carried haplogroup L3b1a+@16124.[16]

At Kilifi, Mtwapa, in Kenya, an individual, dated between 1200 CE and 1450 CE, carried haplogroups E1b1b and L0a2a2a.[16]

At Kilifi, Mtwapa, in Kenya, an individual, dated between 1200 CE and 1450 CE, carried haplogroups E1b1b1b2a1a1a1a1f~ and L0a2a2a.[16]

At Kilifi, Mtwapa, in Kenya, an individual, dated between 1200 CE and 1450 CE, carried haplogroups J1a2a1a2d2b~ and L0a2a2a.[16]

At Kilifi, Mtwapa, in Kenya, an individual, dated between 1200 CE and 1450 CE, carried haplogroup M30d1.[16]

At Kilifi, Mtwapa, in Kenya, an individual, dated between 1200 CE and 1450 CE, carried haplogroup L0a2a2a.[16]

At Kilifi, Mtwapa, in Kenya, an individual, dated between 1200 CE and 1450 CE, carried haplogroup L0a1b1a.[16]

At Kilifi, Mtwapa, in Kenya, an individual, dated between 1226 cal CE and 1297 cal CE, carried haplogroups J1a2a1a2d2b~ and M30d1.[16]

At Kilifi, Mtwapa, in Kenya, an individual, dated between 1323 cal CE and 1423 cal CE, carried haplogroups E1b1b and L0d3.[16]

At Kilifi, Mtwapa, in Kenya, an individual, dated between 1350 CE and 1500 CE, carried haplogroup L1c3a1b.[16]

At Kilifi, Mtwapa, in Kenya, an individual, dated between 1400 CE and 1650 CE, carried haplogroups J1a2a1a2 and L3e3a.[16]

At Kilifi, Mtwapa, in Kenya, an individual, dated between 1408 cal CE and 1442 cal CE, carried haplogroup L0a2a1a2.[16]

At Kilifi, Mtwapa, in Kenya, an individual, dated between 1424 cal CE and 1457 cal CE, carried haplogroup L3a2.[16]

At Kilifi, Mtwapa, in Kenya, an individual, dated between 1435 cal CE and 1469 cal CE, carried haplogroups J and L3d1a.[16]

At Kilifi, Mtwapa, in Kenya, an individual, dated between 1435 cal CE and 1479 cal CE, carried haplogroups J1a2a1a2d2b~ and L0a2a2a.[16]

At Kilifi, Mtwapa, in Kenya, an individual, dated between 1442 cal CE and 1612 cal CE, carried haplogroups J1 and L0a1b1a.[16]

At Kilifi, Mtwapa, in Kenya, an individual, dated between 1445 cal CE and 1609 cal CE, carried haplogroups R1a1a1 and L3b1a1a.[16]

At Kilifi, Mtwapa, in Kenya, an individual, dated between 1446 cal CE and 1611 cal CE, carried haplogroups J1a2a1a2d2b~ and L3d1a1a.[16]

At Kilifi, Mtwapa, in Kenya, an individual, dated between 1446 cal CE and 1614 cal CE, carried haplogroups R1a1a1 and L3e1d1.[16]

At Kilifi, Mtwapa, in Kenya, an individual, dated between 1446 cal CE and 1623 cal CE, carried haplogroups J1a2a1a2d2b~ and L1b1a.[16]

At Kilifi, Mtwapa, in Kenya, an individual, dated between 1450 CE and 1700 CE, carried haplogroups J2a1a1a2a2b and L0a2a1a2.[16]

At Kilifi, Mtwapa, in Kenya, an individual, dated between 1451 cal CE and 1619 cal CE, carried haplogroups J1 and L3e3a.[16]

At Kilifi, Mtwapa, in Kenya, an individual, dated between 1454 cal CE and 1623 cal CE, carried haplogroup L3f1b1a1.[16]

At Kilifi, Mtwapa, in Kenya, an individual, dated between 1457 cal CE and 1626 cal CE, carried haplogroup L2a1b1a.[16]

At Kilifi, Mtwapa, in Kenya, an individual, dated between 1496 cal CE and 1630 cal CE, carried haplogroup L1c3a1b.[16]

At Kilifi, Mtwapa, in Kenya, an individual, dated between 1497 cal CE and 1640 cal CE, carried haplogroup L2a1'2'3'4.[16]

At Kilifi, Mtwapa, in Kenya, an individual, dated between 1508 cal CE and 1654 cal CE, carried haplogroup R0+16189.[16]

Laikipia County

At Kisima Farm/Porcupine Cave, in Laikipia County, Kenya, there were two pastoralists of the Pastoral Neolithic; one carried haplogroups E1b1b1b2b2a1/E-M293 and M1a1, and another carried haplogroup M1a1f.[11][12]

At Kisima Farm/C4, in Laikipia County, Kenya, a pastoralist of the Pastoral Iron Age, carried haplogroups E2 (xE2b)/E-M75 and L3h1a1.[11][12]

At Laikipia District Burial, in Laikipia County, Kenya, a pastoralist of the Pastoral Iron Age carried haplogroup L0a1c1.[11][12]

Lamu

At Lamu, Manda, in Kenya, an individual, dated between 800 CE and 1500 CE, carried haplogroup L3d1a1a.[16]

At Lamu, Manda, in Kenya, an individual, dated between 1400 CE and 1700 CE, carried haplogroups J2 and L2d1a.[16]

At Lamu, Manda, in Kenya, an individual, dated between 1400 CE and 1700 CE, carried haplogroup L2d1a.[16]

At Lamu, Manda, in Kenya, an individual, dated between 1437 cal CE and 1482 cal CE, carried haplogroup L2a1b1.[16]

At Lamu, Manda, in Kenya, an individual, dated between 1456 cal CE and 1621 cal CE, carried haplogroup L2d1a.[16]

At Lamu, Manda, in Kenya, an individual, dated between 1456 cal CE and 1621 cal CE, carried haplogroup J2.[16]

At Lamu, Manda, in Kenya, an individual, dated between 1457 cal CE and 1626 cal CE, carried haplogroups J2b2a2~ and L2d1a.[16]

At Lamu, Manda, in Kenya, an individual, dated between 1485 cal CE and 1629 cal CE, carried haplogroups G2a and L3e3a.[16]

At Lamu, Pate Island, Faza, in Kenya, an individual, dated between 1500 CE and 1700 CE, carried haplogroups E1b1a1a1a2a1a and L3e3a.[16]

Nakuru County

At Prettejohn's Gully, in Nakuru County, Kenya, there were two pastoralists of the early pastoral period; one carried haplogroups E2 (xE2b)/E-M75 and K1a, and another carried haplogroup L3f1b.[11][12]

At Cole's Burial, in Nakuru County, Kenya, a pastoralist of the Pastoral Neolithic carried haplogroups E1b1b1a1a1b1/E-CTS3282 and L3i2.[11][12]

At Rigo Cave, in Nakuru County, Kenya, there were three pastoralists of the Pastoral Neolithic/Elmenteitan, one carried haplogroups E1b1b1b2b2a1/E-M293 and L3f, another carried haplogroups E1b1b1b2b2/E-V1486, likely E-M293, and probably M1a1b, and the last carried haplogroups E1b1b1b2b2a1/E-M293 and L4b2a2c.[11][12]

At Naishi Rockshelter, in Nakuru County, Kenya, there two pastoralists of the Pastoral Neolithic; one carried haplogroups E1b1b1b2b/E-V1515, likely E-M293, and L3x1a, and another carried haplogroups A1b (xA1b1b2a)/A-P108 and L0a2d.[11][12]

At Keringet Cave, in Nakuru County, Kenya, a pastoralist of the Pastoral Neolithic carried haplogroups A1b1b2/A-L427 and L4b2a1, and another pastoralist of the Pastoral Neolithic/Elmenteitan carried haplogroup K1a.[11][12]

At Naivasha Burial Site, in Nakuru County, Kenya, there were five pastoralists of the Pastoral Neolithic; one carried haplogroup L4b2a2b, another carried haplogroups xBT, likely A, and M1a1b, another carried haplogroups E1b1b1b2b2a1/E-M293 and L3h1a1, another carried haplogroups A1b1b2b/A-M13 and L4a1, and the last carried haplogroups E1b1b1b2b2a1/E-M293 and L3x1a.[11][12]

At Njoro River Cave II, in Nakuru County, Kenya, a pastoralist of the Pastoral Neolithic carried haplogroup L3h1a2a1.[11][12]

At Egerton Cave, in Nakuru County, Kenya, a pastoralist of the Pastoral Neolithic/Elmenteitan carried haplogroup L0a1d.[11][12]

At Ilkek Mounds, in Nakuru County, Kenya, a pastoralist of the Pastoral Iron Age carried haplogroups E2 (xE2b)/E-M75 and L0f2a.[11][12]

At Deloraine Farm, in Nakuru County, Kenya, an iron metallurgist of the Iron Age carried haplogroups E1b1a1a1a1a/E-M58 and L5b1.[11][12]

Narok County

At Kasiole 2, in Narok County, Kenya, a pastoralist of the Pastoral Iron Age carried haplogroups E1b1b1b2b/E-V1515, likely E-M293, and L3h1a2a1.[11][12]

At Emurua Ole Polos, in Narok County, Kenya, a pastoralist of the Pastoral Iron Age carried haplogroups E1b1b1b2b2a1/E-M293 and L3h1a2a1.[11][12]

Taita Taveta

At Taita Taveta, Makwasinyi, in Kenya, an individual, dated between 1650 CE and 1950 CE, carried haplogroups E1b1a1a1a2a1a and L4b2a.[16]

At Taita Taveta, Makwasinyi, in Kenya, an individual, dated between 1650 CE and 1950 CE, carried haplogroup L3d1a1a.[16]

At Taita Taveta, Makwasinyi, in Kenya, an individual, dated between 1650 CE and 1950 CE, carried haplogroup L3d1a1a.[16]

At Taita Taveta, Makwasinyi, in Kenya, an individual, dated between 1650 CE and 1950 CE, carried haplogroups E1b1a1a1a2a1a3b1d1c and L1c3b1a.[16]

At Taita Taveta, Makwasinyi, in Kenya, an individual, dated between 1650 CE and 1950 CE, carried haplogroup L3f2a1.[16]

At Taita Taveta, Makwasinyi, in Kenya, an individual, dated between 1650 CE and 1950 CE, carried haplogroup L0f2a.[16]

At Taita Taveta, Makwasinyi, in Kenya, an individual, dated between 1650 CE and 1950 CE, carried haplogroup L5a1a.[16]

At Taita Taveta, Makwasinyi, in Kenya, an individual, dated between 1650 CE and 1950 CE, carried haplogroups E1b1a1a1a2a1a and L2a1+143.[16]

At Taita Taveta, Makwasinyi, in Kenya, an individual, dated between 1650 CE and 1950 CE, carried haplogroup L0d3.[16]

At Taita Taveta, Makwasinyi, in Kenya, an individual, dated between 1650 CE and 1950 CE, carried haplogroup L3e3a.[16]

At Taita Taveta, Makwasinyi, in Kenya, an individual, dated between 1667 cal CE and 1843 cal CE, carried haplogroups E1b1a1a1a2a1a3b1d1c and L2a1+143.[16]

At Taita Taveta, Makwasinyi, in Kenya, an individual, dated between 1698 cal CE and 1950 cal CE, carried haplogroup L0a1a+200.[16]

At Taita Taveta, Makwasinyi, in Kenya, an individual, dated between 1709 cal CE and 1927 cal CE, carried haplogroups E1b1a1a1a2a1a3a1d~ and L3a2.[16]

Tanzania

At Mlambalasi rockshelter, in Tanzania, an individual, dated between 20,345 BP and 17,025 BP, carried undetermined haplogroups.[17]

At Kisese II rockshelter, in Tanzania, an individual, dated between 7240 BP and 6985 BP, carried haplogroups B2b1a~ and L5b2.[17]

At Luxmanda, Tanzania, an individual, estimated to date between 3141 BP and 2890 BP, carried haplogroup L2a1.[15]

At Kuumbi Cave, in Zanzibar, Tanzania, an individual, estimated to date between 1370 BP and 1303 BP, carried haplogroup L4b2a2c.[15]

Karatu District

At Gishimangeda Cave, in Karatu District, Tanzania, there were eleven pastoralists of the Pastoral Neolithic; one carried haplogroups E1b1b1a1b2/E-V22 and HV1b1, another carried haplogroup L0a, another carried haplogroup L3x1, another carried haplogroup L4b2a2b, another carried haplogroups E1b1b1b2b2a1/E-M293 and L3i2, another carried haplogroup L3h1a2a1, another carried haplogroups E1b1b1b2b2/E-V1486, likely E-M293 and L0f2a1, and another carried haplogroups E1b1b1b2b2/E-V1486, likely E-M293, and T2+150; while most of the haplogroups among three pastoralists went undetermined, one was determined to carry haplogroup BT, likely B.[11][12]

Kilwa Coast

At Kilwa, Coast, in Tanzania, an individual, dated between 1300 CE and 1600 CE, carried haplogroups J2a2a1a1a2a~ and L2a1h.[16]

At Kilwa, Coast, in Tanzania, an individual, dated between 1300 CE and 1600 CE, carried haplogroup L3b1a11.[16]

Lindi

At Lindi, in Tanzania, an individual, dated between 1511 cal CE and 1664 cal CE, carried haplogroups E1b1a1a1a2a1a3a1d~ and L0a1a2.[16]

Pemba Island

At Makangale Cave, on Pemba Island, Tanzania, an individual, estimated to date between 1421 BP and 1307 BP, carried haplogroup L0a.[15]

At Makangale Cave, on Pemba Island, Tanzania, an individual, estimated to date between 639 BP and 544 BP, carried haplogroup L2a1a2.[15]

Songo Mnara

At Songo Mnara, in Tanzania, an individual, dated between 1294 cal CE and 1392 cal CE, carried haplogroups R1a and L3e3a.[16]

At Songo Mnara, in Tanzania, an individual, dated between 1402 cal CE and 1437 cal CE, carried haplogroup L3e2b1a2.[16]

At Songo Mnara, in Tanzania, an individual, dated between 1412 cal CE and 1446 cal CE, carried haplogroup L3d1a1a.[16]

At Songo Mnara, in Tanzania, an individual, dated between 1418 cal CE and 1450 cal CE, carried haplogroups E1b1a1~ and L3e2b.[16]

At Songo Mnara, in Tanzania, an individual, dated between 1508 cal CE and 1648 cal CE, carried haplogroup L3d1a1a1.[16]

At Songo Mnara, in Tanzania, an individual, dated between 1516 cal CE and 1667 cal CE, carried haplogroups E1b1b1b2b2a1a~ and L3a1b.[16]

At Songo Mnara, in Tanzania, an individual, dated between 1629 cal CE and 1794 cal CE, carried haplogroups E1b1b1a1a1b2~ and L3d1a1a1.[16]

Uganda

At Munsa, in Uganda, an individual, dated to the Later Iron Age (500 BP), carried haplogroup L3b1a1.[13][14]

Y-Chromosomal DNA

Further information: Y-DNA haplogroups in populations of Sub-Saharan Africa

As of 19,000 years ago, Africans, bearing haplogroup E1b1a-V38, likely traversed across the Sahara, from east to west.[18]

Before the slave trade period, East Africans, who carried haplogroup E1b1a-M2, expanded into Arabia, resulting in various rates of inheritance throughout Arabia (e.g., 2.8% Qatar, 3.2% Yemen, 5.5% United Arab Emirates, 7.4% Oman).[19]

Mitochondrial DNA

Further information: Macro-haplogroup L (mtDNA), Haplogroup M (mtDNA), and Haplogroup N (mtDNA)

In 150,000 BP, Africans (e.g., Central Africans, East Africans) bearing haplogroup L1 diverged.[20] In 130,000 BP, Africans bearing haplogroup L5 diverged in East Africa.[20] Between 130,000 BP and 75,000 BP, behavioral modernity emerged among Southern Africans and long-term interactions between the regions of Southern Africa and Eastern Africa became established.[20] Between 75,000 BP and 60,000 BP, Africans bearing haplogroup L3 emerged in East Africa and eventually migrated into and became present in modern West Africans, Central Africans, and non-Africans.[20] Amid the Holocene, including the Holocene Climate Optimum in 8000 BP, Africans bearing haplogroup L2 spread within West Africa and Africans bearing haplogroup L3 spread within East Africa.[20] As the largest migration since the Out of Africa migration, migration from Sub-Saharan Africa toward the North Africa occurred, by West Africans, Central Africans, and East Africans, resulting in migrations into Europe and Asia; consequently, Sub-Saharan African mitochondrial DNA was introduced into Europe and Asia.[20] During the early period of the Holocene, 50% of Sub-Saharan African mitochondrial DNA was introduced into North Africa by West Africans and the other 50% was introduced by East Africans.[20] During the modern period, a greater number of West Africans introduced Sub-Saharan African mitochondrial DNA into North Africa than East Africans.[20] Between 15,000 BP and 7000 BP, 86% of Sub-Saharan African mitochondrial DNA was introduced into Southwest Asia by East Africans, largely in the region of Arabia, which constitute 50% of Sub-Saharan African mitochondrial DNA in modern Southwest Asia.[20] In the modern period, 68% of Sub-Saharan African mitochondrial DNA was introduced by East Africans and 22% was introduced by West Africans, which constitutes 50% of Sub-Saharan African mitochondrial DNA in modern Southwest Asia.[20]

Autosomal DNA

From the region of Kenya and Tanzania to South Africa, eastern Bantu-speaking Africans constitute a north to south genetic cline; additionally, from eastern Africa to toward southern Africa, evidence of genetic homogeneity is indicative of a serial founder effect and admixture events having occurred between Bantu-speaking Africans and other African populations by the time the Bantu migration had spanned into South Africa.[21]

Across all areas of Madagascar, the average ancestry for the Malagasy people was found to be 4% West Eurasian, 37% Austronesian, and 59% Bantu.[22]

Medical DNA

The genomes of Africans commonly found to undergo adaptation are regulatory DNA, and many cases of adaptation found among Africans relate to diet, physiology, and evolutionary pressures from pathogens.[21] Throughout Sub-Saharan Africa, genetic adaptation (e.g., rs334 mutation, Duffy blood group, increased rates of G6PD deficiency, sickle cell disease) to malaria has been found among Sub-Saharan Africans, which may have initially developed in 7300 BP.[21] Sub-Saharan Africans have more than 90% of the Duffy-null genotype.[23] In the highlands of Ethiopia, genetic adaptation (e.g., rs10803083, an SNP associated with the rate and function of hemoglobin; BHLHE41, a gene associated with circadian rhythm and hypoxia response; EGNL1, a gene strongly associated with oxygen homeostasis in mammals) to hypoxia and low atmospheric pressure has been found among the Amhara people, which may have developed within the past 5000 years.[21] In Tanzania, genetic adaptation (e.g., greater amount of amylase genes than in African populations that consume low-starch foods) has been found in the Hadza people due to a food diet that especially includes consumption of tubers.[21]

References

  1. ^ Osborne, Anne H.; et al. (October 2008). "A humid corridor across the Sahara for the migration of early modern humans out of Africa 120,000 years ago". Proceedings of the National Academy of Sciences of the United States of America. 105 (43): 16444–16447. Bibcode:2008PNAS..10516444O. doi:10.1073/pnas.0804472105. PMC 2575439. PMID 18936490. S2CID 10418009.
  2. ^ Drake, Nick; Breeze, Paul (2016). "Climate Change and Modern Human Occupation of the Sahara from MIS 6-2". Africa from MIS 6-2. Vertebrate Paleobiology and Paleoanthropology. Africa from MIS 6-2. pp. 103–122. doi:10.1007/978-94-017-7520-5_6. ISBN 978-94-017-7519-9. S2CID 131383927.
  3. ^ El-Shenawy, Mohammed I.; et al. (2018). "Speleothem evidence for the greening of the Sahara and its implications for the early human dispersal out of sub-Saharan Africa". Quaternary Science Reviews. 188: 67–76. Bibcode:2018QSRv..188...67E. doi:10.1016/j.quascirev.2018.03.016. S2CID 134694090.
  4. ^ Scheele, Judith (Aug 2016). Crossroads Regions: The Sahara. Oxford Handbooks Online. doi:10.1093/oxfordhb/9780199935369.013.18. ISBN 978-0-19-993536-9.
  5. ^ Wippel, Steffen (2020). "The Sahara as a Bridge, Not a Barrier: An Essay and Book Review on Recent Transregional Perspectives". Neue Politische Literatur. 65 (3): 449–472. doi:10.1007/s42520-020-00318-y. S2CID 224855920.
  6. ^ Bergström A, Stringer C, Hajdinjak M, Scerri EM, Skoglund P (February 2021). "Origins of modern human ancestry". Nature. 590 (7845): 229–237. Bibcode:2021Natur.590..229B. doi:10.1038/s41586-021-03244-5. PMID 33568824. S2CID 231883210.
  7. ^ Llorente MG, Jones ER, Eriksson A, Siska V, Arthur KW, Arthur JW, et al. (November 2015). "Ancient Ethiopian genome reveals extensive Eurasian admixture throughout the African continent". Science. 350 (6262): 820–822. Bibcode:2015Sci...350..820L. doi:10.1126/science.aad2879. hdl:2318/1661894. PMID 26449472. S2CID 25743789.
  8. ^ Llorente MG, Jones ER, Eriksson A, Siska V, Arthur KW, Arthur JW, et al. (13 November 2015). "Supplementary Materials for Ancient Ethiopian genome reveals extensive Eurasian admixture in Eastern Africa". Science. 350 (6262): 820–822. Bibcode:2015Sci...350..820L. doi:10.1126/science.aad2879. hdl:2318/1661894. PMID 26449472. S2CID 25743789.
  9. ^ Hellenthal G, Bird N, Morris S (April 2021). "Structure and ancestry patterns of Ethiopians in genome-wide autosomal DNA". Human Molecular Genetics. 30 (R1): R42 – R48. doi:10.1093/hmg/ddab019. PMC 8242491. PMID 33547782.
  10. ^ Pagani, Luca; Kivisild, Toomas; Tarekegn, Ayele; Ekong, Rosemary; Plaster, Chris; Gallego Romero, Irene; Ayub, Qasim; Mehdi, S. Qasim; Thomas, Mark G.; Luiselli, Donata; Bekele, Endashaw (2012-07-13). "Ethiopian genetic diversity reveals linguistic stratification and complex influences on the Ethiopian gene pool". American Journal of Human Genetics. 91 (1): 83–96. doi:10.1016/j.ajhg.2012.05.015. ISSN 1537-6605. PMC 3397267. PMID 22726845.
  11. ^ a b c d e f g h i j k l m n o p q r s t Prendergast ME, Lipson M, Sawchuk EA, Olalde I, Ogola CA, Rohland N, et al. (July 2019). "Ancient DNA reveals a multistep spread of the first herders into sub-Saharan Africa". Science. 365 (6448): 6275. Bibcode:2019Sci...365.6275P. doi:10.1126/science.aaw6275. PMC 6827346. PMID 31147405.
  12. ^ a b c d e f g h i j k l m n o p q r s t Prendergast ME, Lipson M, Sawchuk EA, Olalde I, Ogola CA, Rohland N, et al. (5 July 2019). "Supplementary Materials for Ancient DNA reveals a multistep spread of the first herders into sub-Saharan Africa". Science. 365 (6448): eaaw6275. Bibcode:2019Sci...365.6275P. doi:10.1126/science.aaw6275. PMC 6827346. PMID 31147405.
  13. ^ a b c d e f Wang K, Goldstein S, Bleasdale M, Clist B, Bostoen K, Bakwa-Lufu P, et al. (June 2020). "Ancient genomes reveal complex patterns of population movement, interaction, and replacement in sub-Saharan Africa". Science Advances. 6 (24): eaaz0183. Bibcode:2020SciA....6..183W. doi:10.1126/sciadv.aaz0183. PMC 7292641. PMID 32582847.
  14. ^ a b c d e f Wang K, Goldstein S, Bleasdale M, Clist B, Bostoen K, Bakwa-Lufu P, et al. (June 2020). "Ancient genomes reveal complex patterns of population movement, interaction, and replacement in sub-Saharan Africa". Science Advances. 6 (24): eaaz0183. Bibcode:2020SciA....6..183W. doi:10.1126/sciadv.aaz0183. PMC 7292641. PMID 32582847.
  15. ^ a b c d e Skoglund P, Thompson JC, Prendergast ME, Mittnik A, Sirak K, Hajdinjak M, et al. (September 2017). "Reconstructing Prehistoric African Population Structure". Cell. 171 (1): 59–71.e21. doi:10.1016/j.cell.2017.08.049. PMC 5679310. PMID 28938123.
  16. ^ a b c d e f g h i j k l m n o p q r s t u v w x y z aa ab ac ad ae af ag ah ai aj ak al am an ao ap aq ar as at au av aw ax ay az ba bb bc bd be bf bg bh bi bj bk bl bm bn bo bp bq br bs bt bu bv bw Brielle, Esther S.; et al. (March 29, 2023). "Supplementary Data Files for Entwined African and Asian genetic roots of medieval peoples of the Swahili coast". Nature. 615 (7954): 866–873. Bibcode:2023Natur.615..866B. doi:10.1038/s41586-023-05754-w. ISSN 0028-0836. OCLC 9819552636. PMC 10060156. PMID 36991187. S2CID 250534036.
  17. ^ a b Lipson, Mark; et al. (23 February 2022). "Extended Data Table 1 Ancient individuals analysed in this study: Ancient DNA and deep population structure in sub-Saharan African foragers". Nature. 603 (7900): 290–296. Bibcode:2022Natur.603..290L. doi:10.1038/s41586-022-04430-9. ISSN 0028-0836. OCLC 9437356581. PMC 8907066. PMID 35197631. S2CID 247083477.
  18. ^ Shriner, Daniel; Rotimi, Charles N. (April 2018). "Whole-Genome-Sequence-Based Haplotypes Reveal Single Origin of the Sickle Allele during the Holocene Wet Phase". American Journal of Human Genetics. 102 (4): 547–556. doi:10.1016/j.ajhg.2018.02.003. ISSN 0002-9297. OCLC 7353789016. PMC 5985360. PMID 29526279.
  19. ^ Neves da Nova Fernandes VC. "High-resolution characterization of genetic markers in the Arabian Peninsula and Near East" (PDF). White Rose eTheses Online. University of Leeds.
  20. ^ a b c d e f g h i j Sá, Luísa; et al. (16 August 2022). "Phylogeography of Sub-Saharan Mitochondrial Lineages Outside Africa Highlights the Roles of the Holocene Climate Changes and the Atlantic Slave Trade". International Journal of Molecular Sciences. 23 (16): 9219. doi:10.3390/ijms23169219. ISSN 1661-6596. OCLC 9627558751. PMC 9408831. PMID 36012483. S2CID 251653686.
  21. ^ a b c d e Pfennig, Aaron; et al. (March 29, 2023). "Evolutionary Genetics and Admixture in African Populations". Genome Biology and Evolution. 15 (4): evad054. doi:10.1093/gbe/evad054. OCLC 9817135458. PMC 10118306. PMID 36987563. S2CID 257803764.
  22. ^ Heiske M, Alva O, Pereda-Loth V, Van Schalkwyk M, Radimilahy C, Letellier T, et al. (April 2021). "Genetic evidence and historical theories of the Asian and African origins of the present Malagasy population". Human Molecular Genetics. 30 (R1): R72 – R78. doi:10.1093/hmg/ddab018. PMID 33481023.
  23. ^ Wonkam, Ambroise; Adeyemo, Adebowale (March 8, 2023). "Leveraging our common African origins to understand human evolution and health" (PDF). Cell Genomics. 3 (3): 100278. doi:10.1016/j.xgen.2023.100278. PMC 10025516. PMID 36950382. S2CID 257458855.

Read other articles:

Askar Perang Sabil

Monumen Askar Perang Sabil di Kampung Kauman, Yogyakarta. Askar Perang Sabil adalah sebuah organisasi kelaskaran yang terbentuk atas prakarsa para ulama Yogyakarta yang sebagian besar berasal dari kalangan Muhammadiyah. APS dapat dikatakan sebagai organisasi kelaskaran bentukan Muhammadiyah.[1][2][3][4][5][6][7] Asal-usul APS terbentuk dari keprihatinan para ulama Muhammadiyah yang melihat bahwa kondisi bangsa Indonesia kian mengkhawatir...

 

Fahd dari Arab Saudi

Penjaga Dua Kota Suci Fahd dari Arab SaudiNama dalam bahasa asli(ar) فهد بن عبد العزيز آل سعود BiografiKelahiran1921 Riyadh Kematian1r Agustus 2005 (83/84 tahun)Riyadh Penyebab kematianRadang paru-paru Tempat pemakamanPemakaman Al Oud Riyadh   Raja Arab Saudi 13 Juni 1982 – 1r Agustus 2005 ← Khalid dari Arab Saudi – Abdullah dari Arab Saudi →   Perdana Menteri Arab Saudi 13 Juni 1982 – 1r Agustus 2005 ...

 

Tenggelamnya KMP Tampomas II

Musibah KMP Tampomas IIFoto dari helikopter yang menunjukkan terbakarnya KMP Tampomas di Laut Jawa.RincianTanggal27 Januari 1981; 43 tahun lalu (1981-01-27)Waktu12.45 WIBLetakKepulauan Masalembo, Laut Jawa275 km (171 mi) dari Surabaya, Jawa TimurKoordinatKoordinat: 5°18′00″S 114°15′22″E / 5.300°S 114.2560°E / -5.300; 114.2560NegaraIndonesiaJalurPadang-Jakarta-Ujung PandangOperatorPT PelniPemilikPemerintah IndonesiaLayananTransportasi umum Kap...

The 2000's Greatest

Artikel ini tidak memiliki referensi atau sumber tepercaya sehingga isinya tidak bisa dipastikan. Tolong bantu perbaiki artikel ini dengan menambahkan referensi yang layak. Tulisan tanpa sumber dapat dipertanyakan dan dihapus sewaktu-waktu.Cari sumber: The 2000's Greatest – berita · surat kabar · buku · cendekiawan · JSTOR The 2000's GreatestAlbum kompilasi karya Dewa 19Dirilis2016Direkam2007–2011GenreRock, Hard Rock, Pop rock, Rock alternatifDur...

 

العلاقات الأسترالية البالاوية

العلاقات الأسترالية البالاوية أستراليا بالاو   أستراليا   بالاو تعديل مصدري - تعديل   العلاقات الأسترالية البالاوية هي العلاقات الثنائية التي تجمع بين أستراليا وبالاو.[1][2][3][4][5] مقارنة بين البلدين هذه مقارنة عامة ومرجعية للدولتين: وجه الم...

 

GTx-758

Chemical compound GTx-758Clinical dataOther namesCapesarisRoutes ofadministrationBy mouthLegal statusLegal status US: Investigational New Drug Identifiers IUPAC name 3-fluoro-N-(4-fluorophenyl)-4-hydroxy-N-(4-hydroxyphenyl)benzamide CAS Number938067-78-8PubChem CID46861578ChemSpider34954411UNII406GR1485ZChEMBLChEMBL3545017Chemical and physical dataFormulaC19H13F2NO3Molar mass341.314 g·mol−13D model (JSmol)Interactive image SMILES C1=CC(=CC=C1N(C2=CC=C(C=C2)F)C(=O)C3=CC(=C(C=C3)O)...

趙家驤

追晉陸軍二級上將趙家驤將軍个人资料出生1910年 大清河南省衛輝府汲縣逝世1958年8月23日(1958歲—08—23)（47—48歲） † 中華民國福建省金門縣国籍 中華民國政党 中國國民黨获奖 青天白日勳章（追贈）军事背景效忠 中華民國服役 國民革命軍 中華民國陸軍服役时间1924年－1958年军衔 二級上將 （追晉）部队四十七師指挥東北剿匪總司令部參謀長陸軍�...

 

Claas

Global agricultural machinery manufacturer CLAAS KGaA mbHCompany typeGmbH & Co. KGaAIndustryAgricultural machineryFounded1913; 111 years ago (1913)FounderAugust ClaasHeadquartersHarsewinkel, GermanyArea servedWorldwideKey peopleCEO Thomas BöckProductsTractorsCombinesDiesel enginesGearboxes and AxlesBalersForage harvestersTelescopic handlersWheel loadersFarming Information TechnologyRevenue €3.898 bn (2019)[1]Number of employees11,448 (2019)Websitewww.claas.com...

 

大字 (数字)

この項目には、一部のコンピュータや閲覧ソフトで表示できない文字が含まれています（詳細）。 数字の大字（だいじ）は、漢数字の一種。通常用いる単純な字形の漢数字（小字）の代わりに同じ音の別の漢字を用いるものである。 概要 壱万円日本銀行券（「壱」が大字） 弐千円日本銀行券（「弐」が大字） 漢数字には「一」「二」「三」と続く小字と、「壱」「�...

Coach Design

Australian bus bodybuilder Coach DesignAustralia Wide Coaches Scania K480EB in Wagga Wagga in November 2009IndustryBus manufacturingPredecessorGamer Brewer & WinterHeadquartersArcherfieldParentChryss JamiesonWebsitecoachdesign.com.au Coach Design is an Australian bus bodybuilder in Archerfield, Brisbane.[1] Until 1992 it traded as GBW.[2] References ^ Our Products Coach Design ^ Bodybuilders Fleetline issue 239 November 1995 page 195 External links Media related to Coach D...

 

رجال لا يعرفون الحب (فيلم)

رجال لا يعرفون الحب رجال لا يعرفون الحب  الصنف دراما، جريمة تاريخ الصدور 16 يوليو 1979 مدة العرض 100 دقيقة البلد  مصر اللغة الأصلية العربية الطاقم المخرج يحيى العلمي الإنتاج چيبكو (تاكفور أنطونيان) الكاتب محمد عثمان البطولة فريد شوقيحسين فهميميرفت أمين صناعة سينمائية تصو...

 

Kish civilization

An ancient mound at the city of Kish, Mesopotamia, Babel Governorate, Iraq According to a theory proposed by Ignace Gelb, the Kish civilization encompassed the sites of Ebla and Mari in the Levant, Nagar in the north,[1] and the proto-Akkadian sites of Abu Salabikh and Kish in central Mesopotamia[2][3][better source needed] in to the early East Semitic era in Mesopotamia and the Levant. The epoch began in the early 4th millennium BC and ended with the...

Coyote (racial category)

Racial term De Mestizo y de India; Coyote. Miguel Cabrera, 1763, oil on canvas, Waldo-Dentzel Art Center. De mestizo e india, sale coiote. Anonymous, 18th century (From a Mestizo man and an Amerindian woman, a coyote is begotten). De Castizo y India, Coyota. Anonymous, 18th century Mexico. Coyote (fem. Coyota) (from the Nahuatl word coyotl, coyote) is a colonial Spanish American racial term for a mixed-race person casta that usually refers to a person born of parents, one of whom a Mestizo (m...

 

Monsigliolo

Questa voce o sezione sull'argomento centri abitati della Toscana non cita le fonti necessarie o quelle presenti sono insufficienti. Puoi migliorare questa voce aggiungendo citazioni da fonti attendibili secondo le linee guida sull'uso delle fonti. MonsigliolofrazioneMonsigliolo – Veduta LocalizzazioneStato Italia Regione Toscana Provincia Arezzo Comune Cortona TerritorioCoordinate43°14′58″N 11°56′36″E43°14′58″N, 11°56′36″E (Monsigliolo) Abitant...

 

Arthur Conan Doyle

British writer and physician (1859–1930) Conan Doyle redirects here. For the rugby player, see Conan Doyle (rugby union). For the South African cricketer, see Conan Doyle (cricketer). SirArthur Conan DoyleKStJ, DLDoyle in 1914BornArthur Ignatius Conan Doyle(1859-05-22)22 May 1859Edinburgh, ScotlandDied7 July 1930(1930-07-07) (aged 71)Crowborough, Sussex, EnglandOccupation Writer physician EducationUniversity of EdinburghGenre Detective fiction fantasy science fiction historical...

رحل

  لمعانٍ أخرى، طالع رحل (توضيح). رعاة رحل في التبت الرُّحّل لفظ يطلق بصورة عامة على المجموعات البشرية التي يعتمد نمط معيشتها على الترحال والتنقل بدلاً من العيش في القرى والمدن.[1] ويصنف الرحل إلى عدة أنواع من الناحية الاقتصادية (مثل الرعاة الرحل)، ويصنفون إلى عدة أنو...

 

Senapan lontak

Senapan lontak atau musket Senapan lontak atau senapan lantak yang juga dikenal di luar negeri dengan nama musket adalah senjata yang populer di antara abad ke-15 sampai pertengahan abad ke-19. Senapan lontak hanya dapat ditembakkan sekali saja, setelah diisi dengan amunisi bola timah dan mesiu, dan diisi dari depan moncong laras senapan. Etimologi dan sejarah Istilah senapan lontak berasal dari kata senapan dan lantak. Kata senapan adalah korupsi dari kata Belanda snappaan.[1] Lantak...

 

Alexandria Ocasio-Cortez

American politician (born 1989) AOC redirects here. For other uses, see AOC (disambiguation). Alexandria Ocasio-CortezOfficial portrait, 2019Member of the U.S. House of Representativesfrom New York's 14th districtIncumbentAssumed office January 3, 2019Preceded byJoe Crowley Personal detailsBorn (1989-10-13) October 13, 1989 (age 34)New York City, U.S.Political partyDemocraticOther politicalaffiliationsWorking Families Party[a]Democratic Socialists of America[b...

Sex Machines Museum

Sex museum in Prague Entrance to the museum Three BDSM masks on display in the Sex Machines Museum. Sex Machines Museum (abbreviated SMM[1]) is a sex museum in Prague, Czech Republic,[2] which has a collection of sex devices.[3] Established in 2002, it is located near the Old Town Square.[4] The official website of the Sex Machines Museum describes itself as an exposition of mechanical erotic appliances, the purpose of which is to bring pleasure and allow extra...

 

Prototype Verification System

Pour les articles homonymes, voir PVS. Cette page est une très courte ébauche oubliée. À l'instar du célèbre « Une pomme est un fruit » qui a marqué l'histoire de Wikipédia, n'hésitez pas à la développer (comment ?) Vous pouvez également enrichir les pages proposées dans la rubrique « Pommes à croquer » du bistro du jour. Cet article est une ébauche concernant l’informatique. Vous pouvez partager vos connaissances en l’améliorant (comment ...