Old page wikitext, before the edit (old_wikitext) | '[[File:ADMIXTURE analysis of Horn of Africa populations in a broad context.png|thumb|Population structure of African populations in a broad context. ADMIXTURE analysis of 2,194 individuals from 81 populations for 16,420 SNPs reveals both well-established and novel ancestry components in African populations.]]
The '''genetic history of Africa''' is composed of the overall [[genetic history]] of [[Africa|African populations]], including the regional genetic histories of [[North Africa]], [[West Africa]], [[East Africa]], [[Central Africa]], and [[Southern Africa]], as well as the [[recent African origin of modern humans|recent origin of modern humans in Africa]]. The [[Sahara]] served as a trans-regional passageway and place of dwelling for people in Africa during various [[Interstadial|humid phases]]<ref name="Osborn">{{cite journal |vauthors=Osborne AH, Vance D, Rohling EJ, Barton N, Rogerson M, Fello N |title=A humid corridor across the Sahara for the migration of early modern humans out of Africa 120,000 years ago |journal=Proceedings of the National Academy of Sciences of the United States of America |volume=105 |issue=43 |pages=16444–16447 |date=October 2008 |pmid=18936490 |pmc=2575439 |doi=10.1073/pnas.0804472105 |s2cid=10418009 |doi-access=free |bibcode=2008PNAS..10516444O}}</ref><ref name="Drake">{{cite book |vauthors=Drake N, Breeze P |title=Africa from MIS 6-2 |chapter=Climate Change and Modern Human Occupation of the Sahara from MIS 6-2 |chapter-url=https://link.springer.com/chapter/10.1007/978-94-017-7520-5_6 |website=SpringerLink |series=Vertebrate Paleobiology and Paleoanthropology |year=2016 |pages=103–122 |publisher=Africa from MIS 6-2 |doi=10.1007/978-94-017-7520-5_6 |isbn=978-94-017-7519-9}}</ref><ref name="El-Shenawy">{{cite journal |vauthors=El-Shenawy MI, Kim ST, Schwarcz HP, Asmerom Y, Polyak VJ |title=Speleothem evidence for the greening of the Sahara and its implications for the early human dispersal out of sub-Saharan Africa |url=https://www.sciencedirect.com/science/article/abs/pii/S0277379117307436 |journal=Quaternary Science Reviews |year=2018 |volume=188 |pages=67–76 |doi=10.1016/j.quascirev.2018.03.016 |bibcode=2018QSRv..188...67E}}</ref> and periods throughout the [[history of Africa]].<ref name="Scheele">{{cite journal |vauthors=Scheele J |title=Crossroads Regions: The Sahara |url=https://www.academia.edu/37787505 |website=Academia |publisher=Oxford Handbooks Online}}</ref><ref name="Wippel">{{cite journal |vauthors=Wippel S |title=The Sahara as a Bridge, Not a Barrier: An Essay and Book Review on Recent Transregional Perspectives |journal=Neue Politische Literatur |year=2020 |volume=65 |issue=3 |pages=449–472 |doi=10.1007/s42520-020-00318-y |doi-access=free}}</ref>
==Overview==
[[File:PCA of sub-Saharan Africa populations.png|thumb|(A) the origin of the 46 African ethnic groups used in the analysis; ethnic groups from similar regions are given the same colour, but different shapes. (B) PCA shows that the first major axis of variation in Africa (PC1, y-axis) splits southern groups from the rest of Africa, each symbol represents an individual; PC2 (x-axis) reflects ethno-linguistic differences, with Niger-Congo and Nilo-Saharan speakers split from Afroasiatic speakers. (C) The third principle component (PC3, x-axis) represents geographical separation of Niger-Congo speakers, forming a cline from west to east Africans.]]
The people of [[Africa]] are characterized by regional genetic substructure and heterogeneity, depending on the respective ethno-linguistic identity, and, in part, explainable by the "multiregional evolution" of modern human lineages in various multiple regions of the African continent, as well as later admixture events, including back-migrations from Eurasia, of both highly differentiated West- and East-Eurasian components.<ref name="The deep population history in Afri">{{cite journal |vauthors=Hollfelder N, Breton G, Sjödin P, Jakobsson M |title=The deep population history in Africa |journal=Human Molecular Genetics |volume=30 |issue=R1 |pages=R2–R10 |date=April 2021 |pmid=33438014 |pmc=8117439 |doi=10.1093/hmg/ddab005}}</ref>
Africans' genetic ancestry is largely partitioned by [[geography]] and [[language family]], with populations belonging to the same ethno-linguistic groupings showing high genetic homogeneity and coherence. Gene flow, consistent with both short- and long-range migration events followed by extensive admixture and [[bottleneck effect|bottleneck event]]s, have influenced the regional genetic makeup and demographic structure of Africans. The historical [[Bantu expansion]] had lasting impacts on the modern demographic make up of Africa, resulting in a greater genetic and linguistic homogenization.<ref name="Fan 82">{{cite journal |vauthors=Fan S, Kelly DE, Beltrame MH, Hansen ME, Mallick S, Ranciaro A, Hirbo J, Thompson S, Beggs W, Nyambo T, Omar SA, Meskel DW, Belay G, Froment A, Patterson N, Reich D, Tishkoff SA |display-authors=6 |title=African evolutionary history inferred from whole genome sequence data of 44 indigenous African populations |journal=Genome Biology |volume=20 |issue=1 |pages=82 |date=April 2019 |pmid=31023338 |pmc=6485071 |doi=10.1186/s13059-019-1679-2}}</ref> Genetic, archeologic, and linguistic studies added extra insight into this movement: "Our results reveal a genetic continuum of Niger–Congo speaker populations across the continent and extend our current understanding of the routes, timing and extent of the Bantu migration."<ref>{{Cite web |title=A great African gene migration |url=https://cosmosmagazine.com/science/a-great-migration-of-genes-in-africa/ |access-date=2022-01-11 |website=cosmosmagazine.com |date=29 October 2020 |language=en-AU}}</ref>
Overall, different African populations display genetic diversity and substructure, but can be clustered in distinct but partially overlapping groupings:<ref name=":5">{{Cite web |last=Ananyo Choudhury, Shaun Aron, Dhriti Sengupta, Scott Hazelhurst, Michèle Ramsay |date=1 August 2018 |title=African genetic diversity provides novel insights into evolutionary history and local adaptations |url=https://academic.oup.com/hmg/article/27/R2/R209/4993963 |access-date=2022-10-21 |website=academic.oup.com}}</ref>
* [[Khoisan]] hunter-gatherer lineages from [[Southern Africa]] represent the deepest lineages, forming a divergent and distinct cluster, shifted away from contemporary "Sub-Saharan Africans", and are as diverged from them as the various Eurasian lineages are. The diverging date of these "Southern hunter-gatherers" from all other human populations is estimated to over 100,000 years ago respectively, with the Khoisan later diverging into two subgroups, northern and southern Khoisan,~30,000 years ago.{{NoteTag|Although the estimated date of this event varies across studies, there is general consensus that the split occurred over 100 thousand years ago (kya) (8,23).}}
* Rain forest foragers such as the [[Baka people (Cameroon and Gabon)|Baka]] and the [[Mbuti people|Mbuti]] diverged from other Sub-Saharan African groups over 60,000 years ago. Eastern groups such as the Mbuti split from Western groups such as the Baka ~20,000 years ago.
* [[Proto-Afroasiatic language|Proto-Afroasiatic]] speakers are suggested to have diverged from other African groups ~50,000 years ago.<ref>{{Cite journal |last=Baker |first=Jennifer L. |last2=Rotimi |first2=Charles N. |last3=Shriner |first3=Daniel |date=2017-05-08 |title=Human ancestry correlates with language and reveals that race is not an objective genomic classifier |url=https://www.nature.com/articles/s41598-017-01837-7 |journal=Scientific Reports |language=en |volume=7 |issue=1 |pages=1572 |doi=10.1038/s41598-017-01837-7 |issn=2045-2322}}</ref>
* [[Niger Congo|Niger-Congo]] and [[Nilo-Saharan languages|Nilo-Saharan]] speakers split around 28,000 years ago.<ref name=":5" />
* [[Austronesian languages|Austronesian-speaking]] [[Malagasy people]] in [[Madagascar]] have received significant [[East Asian people|East/Southeast Asian]] admixture, less among some groups of coastal Southern, Eastern and the Horn of Africa. The estimated date of geneflow is 2,200 years ago.<ref>{{Cite journal |last1=Kusuma |first1=Pradiptajati |last2=Brucato |first2=Nicolas |last3=Cox |first3=Murray P. |last4=Pierron |first4=Denis |last5=Razafindrazaka |first5=Harilanto |last6=Adelaar |first6=Alexander |last7=Sudoyo |first7=Herawati |last8=Letellier |first8=Thierry |last9=Ricaut |first9=François-Xavier |date=2016-05-18 |title=Contrasting Linguistic and Genetic Origins of the Asian Source Populations of Malagasy |journal=Scientific Reports |volume=6 |pages=26066 |doi=10.1038/srep26066 |issn=2045-2322 |pmc=4870696 |pmid=27188237|bibcode=2016NatSR...626066K }}</ref><ref>{{cite journal |vauthors=Cruciani F, Santolamazza P, Shen P, Macaulay V, Moral P, Olckers A, Modiano D, Holmes S, Destro-Bisol G, Coia V, Wallace DC, Oefner PJ, Torroni A, Cavalli-Sforza LL, Scozzari R, Underhill PA |display-authors=6 |title=A back migration from Asia to sub-Saharan Africa is supported by high-resolution analysis of human Y-chromosome haplotypes |journal=American Journal of Human Genetics |volume=70 |issue=5 |pages=1197–1214 |date=May 2002 |pmid=11910562 |pmc=447595 |doi=10.1086/340257}}</ref><ref>{{cite journal |vauthors=Pakstis AJ, Speed WC, Soundararajan U, Rajeevan H, Kidd JR, Li H, Kidd KK |title=Population relationships based on 170 ancestry SNPs from the combined Kidd and Seldin panels |journal=Scientific Reports |volume=9 |issue=1 |pages=18874 |date=December 2019 |pmid=31827153 |pmc=6906462 |doi=10.1038/s41598-019-55175-x |bibcode=2019NatSR...918874P}}</ref><ref name="Fan 82" /><ref name="Identifying and Interpreting Appare">{{cite journal |vauthors=Chen L, Wolf AB, Fu W, Li L, Akey JM |title=Identifying and Interpreting Apparent Neanderthal Ancestry in African Individuals |journal=Cell |volume=180 |issue=4 |pages=677–687.e16 |date=February 2020 |pmid=32004458 |doi=10.1016/j.cell.2020.01.012 |s2cid=210955842}}</ref><ref name=":2">{{Cite journal |last1=van de Loosdrecht |first1=Marieke |last2=Bouzouggar |first2=Abdeljalil |last3=Humphrey |first3=Louise |last4=Posth |first4=Cosimo |last5=Barton |first5=Nick |last6=Aximu-Petri |first6=Ayinuer |last7=Nickel |first7=Birgit |last8=Nagel |first8=Sarah |last9=Talbi |first9=El Hassan |last10=El Hajraoui |first10=Mohammed Abdeljalil |last11=Amzazi |first11=Saaïd |last12=Hublin |first12=Jean-Jacques |last13=Pääbo |first13=Svante |last14=Schiffels |first14=Stephan |last15=Meyer |first15=Matthias |date=2018-05-04 |title=Pleistocene North African genomes link Near Eastern and sub-Saharan African human populations |url=https://www.science.org/doi/10.1126/science.aar8380 |journal=Science |language=en |volume=360 |issue=6388 |pages=548–552 |doi=10.1126/science.aar8380 |pmid=29545507 |bibcode=2018Sci...360..548V |s2cid=206666517 |issn=0036-8075}}</ref>
[[File:Overall genetic position of worldwide populations.jpg|thumb|Geographic location of the samples analyzed in this study (A). PCA of the Khoe-San individuals, Eurasians, West and East Africans before (unmasked, B) and after (masked, C) applying the local ancestry pipeline (146,696 independent SNPs).]]
===Indigenous Africans===
The indigenous population of Africa consists of [[Niger–Congo languages|Niger–Congo speakers]], [[Nilo-Saharan languages|Nilo-Saharan speakers]], the divergent and diverse [[Khoisan languages|Khoisan grouping]], as well as of several unclassified or [[Language isolate|isolated]] ethnolinguistic groupings (see [[:Category:Unclassified languages of Africa|unclassified languages of Africa]]). The origin of the [[Afroasiatic languages]] remains disputed, with some proposing a [[Middle Eastern]] origin, while others support an African origin with varying degrees of Eurasian and African components.<ref>{{Cite journal |last=Baker |first=Jennifer L. |last2=Rotimi |first2=Charles N. |last3=Shriner |first3=Daniel |date=2017-05-08 |title=Human ancestry correlates with language and reveals that race is not an objective genomic classifier |url=https://www.nature.com/articles/s41598-017-01837-7 |journal=Scientific Reports |language=en |volume=7 |issue=1 |pages=1572 |doi=10.1038/s41598-017-01837-7 |issn=2045-2322}}</ref> The [[Austronesian languages]] originated in southern East Asia, and later expanded outgoing from the [[Philippines]].
[[File:Procrustes-transformed PCA plot of genetic variation of worldwide populations.png|thumb|PCA plot of genetic variation of worldwide populations. ('''A''') Geographic coordinates of 53 populations. ('''B''') Procrustes-transformed PCA plot of genetic variation.<ref>{{cite journal |vauthors=Wang C, Zöllner S, Rosenberg NA |title=A quantitative comparison of the similarity between genes and geography in worldwide human populations |journal=PLOS Genetics |volume=8 |issue=8 |pages=e1002886 |date=August 2012 |pmid=22927824 |pmc=3426559 |doi=10.1371/journal.pgen.1002886}}</ref>]]
The [[Niger–Congo languages]] probably originated in or near the area where these languages were spoken prior to [[Bantu expansion]] (i.e. [[West Africa]] or [[Central Africa]]). Its expansion may have been associated with the expansion of agriculture, during in the [[Neolithic|African Neolithic]] period, following the [[African humid period#End|desiccation of the Sahara]] in c. 3500 BCE. [[Proto-Niger-Congo]] may have originated about 10,000 years before present in the "[[Green Sahara]]" of Africa (roughly the [[Sahel]] and southern [[Sahara]]), and that its dispersal can be correlated with the spread of the [[bow and arrow]] by migrating [[hunter-gatherer]]s, which later developed agriculture.<ref>{{Cite journal |vauthors=Manning K, Timpson A |date=2014-10-01 |title=The demographic response to Holocene climate change in the Sahara |journal=Quaternary Science Reviews |language=en |volume=101 |pages=28–35 |doi=10.1016/j.quascirev.2014.07.003 |bibcode=2014QSRv..101...28M |s2cid=54923700 |issn=0277-3791}}</ref><ref name="BlenchAfricanPast">{{Cite book |vauthors=Blench R |title=Archaeology, language, and the African past |publisher=AltaMira Press |year=2006 |isbn=9780759104655}}</ref><ref>{{cite conference |vauthors=Blench R |url=https://llacan.cnrs.fr/nigercongo2/discussions/Can_we_visit_the_graves_of_the_first_Niger.pdf |title=Can we visit the graves of the first Niger-Congo speakers? |conference=2nd International Congress: Towards Proto-Niger-Congo: Comparison and Reconstruction |location=Paris |date=September 2016}}</ref>
Although the validity of the [[Nilo-Saharan languages|Nilo-Saharan family]] remains controversial, the region between [[Chad]], [[Sudan]], and the [[Central African Republic]] is seen as a likely candidate for its homeland prior to its dispersal around 10,000–8,000 BCE.<ref>{{Cite book |vauthors=Vossen R, Dimmendaal GJ |url=https://books.google.com/books?id=wcjXDwAAQBAJ&pg=PA380 |title=The Oxford Handbook of African Languages |date=2020-03-13 |publisher=Oxford University Press |isbn=978-0-19-960989-5 |language=en}}</ref>
The Southern African hunter-gatherers (Khoisan) are suggested to represent the autochthonous hunter-gatherer population of southern Africa, prior to the expansion of Bantu-speakers from Western/Central Africa and East African pastoralists. Khoisan show evidence for Bantu-related admixture, ranging from nearly ~0% to up to ~87.1%.<ref>{{cite journal |vauthors=Vicente M, Jakobsson M, Ebbesen P, Schlebusch CM |title=Genetic Affinities among Southern Africa Hunter-Gatherers and the Impact of Admixing Farmer and Herder Populations |journal=Molecular Biology and Evolution |volume=36 |issue=9 |pages=1849–1861 |date=September 2019 |pmid=31288264 |pmc=6735883 |doi=10.1093/molbev/msz089}}</ref>
===Out-of-Africa event===
[[File:Human migration routes following Out-of-Africa.png|thumb|Human migration routes following Out-of-Africa.]]
[[File:Migraciones humanas en haplogrupos de ADN-Y.PNG|thumb|Most modern Africans display a high level of genetic homogeneity, but contributions from Eurasian populations are substantial, mostly concentrated in the Northeastern part of Africa and Madagascar.]]
The model proposes a "single origin" of ''[[Homo sapiens]]'' in Africa the taxonomic sense. Recent genetic and archeologic data suggests that Homo sapiens-subgroups originated in multiple regions of Africa, not confined to a single region of origin. The ''H. sapiens'' ancestral to proper Eurasians most likely left [[Northeastern Africa]] between 50,000 and 100,000 years ago.<ref name=":2" /> The "recent African origin" model proposes that all modern non-African populations descend from one or several diverse waves of ''H. sapiens'' that left Africa ~70,000 years ago, with some proposing an earlier date.<ref>{{cite journal |vauthors=López S, van Dorp L, Hellenthal G |title=Human Dispersal Out of Africa: A Lasting Debate |journal=Evolutionary Bioinformatics Online |volume=11 |issue=Suppl 2 |pages=57–68 |date=2016-04-21 |pmid=27127403 |pmc=4844272 |doi=10.4137/EBO.S33489}}</ref><ref>{{cite journal |vauthors=Scerri EM, Thomas MG, Manica A, Gunz P, Stock JT, Stringer C, Grove M, Groucutt HS, Timmermann A, Rightmire GP, d'Errico F, Tryon CA, Drake NA, Brooks AS, Dennell RW, Durbin R, Henn BM, Lee-Thorp J, deMenocal P, Petraglia MD, Thompson JC, Scally A, Chikhi L |display-authors=6 |title=Did Our Species Evolve in Subdivided Populations across Africa, and Why Does It Matter? |language=English |journal=Trends in Ecology & Evolution |volume=33 |issue=8 |pages=582–594 |date=August 2018 |pmid=30007846 |pmc=6092560 |doi=10.1016/j.tree.2018.05.005}}</ref><ref>{{Cite web |title=One Species, Many Origins |url=https://www.shh.mpg.de/1474609/pan-african-origins |access-date=2022-01-11 |website=www.shh.mpg.de |language=en}}</ref>
According to a 2020 study by Durvasula et al., there are indications that 2% to 19% (≃6.6 to 7.0%) of the DNA of West African populations may have come from an unknown archaic hominin which split from the ancestor of humans and Neanderthals between 360 kya to 1.02 mya. However, the study also suggests that at least part of this archaic admixture is also present in Eurasians/non-Africans, and that the admixture event or events range from 0 to 124 ka B.P, which includes the period before the Out-of-Africa migration and prior to the African/Eurasian split (thus affecting in part the common ancestors of both Africans and Eurasians/non-Africans).<ref>{{cite journal |vauthors=Durvasula A, Sankararaman S |date=February 2020 |title=Recovering signals of ghost archaic introgression in African populations |journal=Science Advances |volume=6 |issue=7 |pages=eaax5097 |bibcode=2020SciA....6.5097D |doi=10.1126/sciadv.aax5097 |pmc=7015685 |pmid=32095519 |doi-access=free}} "Non-African populations (Han Chinese in Beijing and Utah residents with northern and western European ancestry) also show analogous patterns in the CSFS, suggesting that a component of archaic ancestry was shared before the split of African and non-African populations...One interpretation of the recent time of introgression that we document is that archaic forms persisted in Africa until fairly recently. Alternately, the archaic population could have introgressed earlier into a modern human population, which then subsequently interbred with the ancestors of the populations that we have analyzed here. The models that we have explored here are not mutually exclusive, and it is plausible that the history of African populations includes genetic contributions from multiple divergent populations, as evidenced by the large effective population size associated with the introgressing archaic population...Given the uncertainty in our estimates of the time of introgression, we wondered whether jointly analyzing the CSFS from both the CEU (Utah residents with Northern and Western European ancestry) and YRI genomes could provide additional resolution. Under model C, we simulated introgression before and after the split between African and non-African populations and observed qualitative differences between the two models in the high-frequency–derived allele bins of the CSFS in African and non-African populations (fig. S40). Using ABC to jointly fit the high-frequency–derived allele bins of the CSFS in CEU and YRI (defined as greater than 50% frequency), we find that the lower limit on the 95% credible interval of the introgression time is older than the simulated split between CEU and YRI (2800 versus 2155 generations B.P.), indicating that at least part of the archaic lineages seen in the YRI are also shared with the CEU..."</ref><ref>''[https://www.science.org/doi/10.1126/sciadv.aax5097] {{Webarchive|url=https://web.archive.org/web/20201207165127/https://advances.sciencemag.org/content/advances/suppl/2020/02/10/6.7.eaax5097.DC1/aax5097_SM.pdf|date=7 December 2020}} Supplementary Materials for ''Recovering signals of ghost archaic introgression in African populations", section "S8.2" "We simulated data using the same priors in Section S5.2, but computed the spectrum for both YRI [West African Yoruba] and CEU [a population of European origin] . We found that the best fitting parameters were an archaic split time of 27,000 generations ago (95% HPD: 26,000-28,000), admixture fraction of 0.09 (95% HPD: 0.04-0.17), admixture time of 3,000 generations ago (95% HPD: 2,800-3,400), and an effective population size of 19,700 individuals (95% HPD: 19,300-20,200). We find that the lower bound of the admixture time is further back than the simulated split between CEU and YRI (2155 generations ago), providing some evidence in favor of a pre-Out-of-Africa event. This model suggests that many populations outside of Africa should also contain haplotypes from this introgression event, though detection is difficult because many methods use unadmixed outgroups to detect introgressed haplotypes [Browning et al., 2018, Skov et al., 2018, Durvasula and Sankararaman, 2019] (5, 53, 22). It is also possible that some of these haplotypes were lost during the Out-of-Africa bottleneck."</ref><ref>{{cite journal |vauthors=Durvasula A, Sankararaman S |date=February 2020 |title=Recovering signals of ghost archaic introgression in African populations |journal=Science Advances |volume=6 |issue=7 |pages=eaax5097 |bibcode=2020SciA....6.5097D |doi=10.1126/sciadv.aax5097 |pmc=7015685 |pmid=32095519}}</ref> Another academic paper from 2020 (Chen et al.) found that Africans have higher [[Neanderthal]] ancestry than previously thought. 2,504 African samples from all over Africa were analyzed and tested on Neanderthal ancestry. All African samples showed evidence for minor Neanderthal ancestry, but always at lower levels than observed in Eurasians.<ref name="Identifying and Interpreting Appare2">{{cite journal |vauthors=Chen L, Wolf AB, Fu W, Li L, Akey JM |date=February 2020 |title=Identifying and Interpreting Apparent Neanderthal Ancestry in African Individuals |journal=Cell |volume=180 |issue=4 |pages=677–687.e16 |doi=10.1016/j.cell.2020.01.012 |pmid=32004458 |s2cid=210955842}}</ref>
===Geneflow between Eurasian and African populations===
{{See also|Eurasian backflow}}
[[File:Pre-Neolithic and Neolithic migration events in Africa (excluding Basal-West-Eurasian geneflow during the Paleolithic).jpg|thumb|Pre-Neolithic and Neolithic migration events in Africa.<ref>{{Cite journal |last=Vicente |first=Mário |last2=Schlebusch |first2=Carina M |date=2020-06-01 |title=African population history: an ancient DNA perspective |url=https://www.sciencedirect.com/science/article/pii/S0959437X20300599 |journal=Current Opinion in Genetics & Development |series=Genetics of Human Origin |language=en |volume=62 |pages=8–15 |doi=10.1016/j.gde.2020.05.008 |issn=0959-437X}}</ref>]]
Significant Eurasian admixture is found in [[Northern Africa]], and among specific ethnic groups of the [[Horn of Africa]], as well as among the [[Malagasy people]] of [[Madagascar]]. Various genome studies found evidence for multiple [[Prehistory|prehistoric]] back-migrations from various [[Eurasia]]n populations and subsequent admixture with native groups.<ref>{{cite journal |vauthors=Busby GB, Band G, Si Le Q, Jallow M, Bougama E, Mangano VD, Amenga-Etego LN, Enimil A, Apinjoh T, Ndila CM, Manjurano A, Nyirongo V, Doumba O, Rockett KA, Kwiatkowski DP, Spencer CC |display-authors=6 |title=Admixture into and within sub-Saharan Africa |journal=eLife |volume=5 |date=June 2016 |pmid=27324836 |pmc=4915815 |doi=10.7554/eLife.15266}}</ref> West-Eurasian geneflow arrived to Northern Africa during the Paleolithic (30,000 to 15,000 years ago), followed by other pre-Neolithic and Neolithic migration events. Genetic data on the [[Taforalt]] samples "demonstrated that Northern Africa received significant amounts of gene-flow from Eurasia predating the Holocene and development of farming practices". Medieval geneflow events, such as the [[Arab expansion]] also left traces in various African populations.<ref>{{Cite journal |last=van de Loosdrecht |first=Marieke |last2=Bouzouggar |first2=Abdeljalil |last3=Humphrey |first3=Louise |last4=Posth |first4=Cosimo |last5=Barton |first5=Nick |last6=Aximu-Petri |first6=Ayinuer |last7=Nickel |first7=Birgit |last8=Nagel |first8=Sarah |last9=Talbi |first9=El Hassan |last10=El Hajraoui |first10=Mohammed Abdeljalil |last11=Amzazi |first11=Saaïd |last12=Hublin |first12=Jean-Jacques |last13=Pääbo |first13=Svante |last14=Schiffels |first14=Stephan |last15=Meyer |first15=Matthias |date=2018-05-04 |title=Pleistocene North African genomes link Near Eastern and sub-Saharan African human populations |url=https://www.science.org/doi/10.1126/science.aar8380 |journal=Science |language=en |volume=360 |issue=6388 |pages=548–552 |doi=10.1126/science.aar8380 |issn=0036-8075}}</ref><ref>{{Cite journal |last=Vicente |first=Mário |last2=Schlebusch |first2=Carina M |date=2020-06-01 |title=African population history: an ancient DNA perspective |url=https://www.sciencedirect.com/science/article/pii/S0959437X20300599 |journal=Current Opinion in Genetics & Development |series=Genetics of Human Origin |language=en |volume=62 |pages=8–15 |doi=10.1016/j.gde.2020.05.008 |issn=0959-437X}}</ref><ref>{{Cite journal |last=Serra-Vidal |first=Gerard |last2=Lucas-Sanchez |first2=Marcel |last3=Fadhlaoui-Zid |first3=Karima |last4=Bekada |first4=Asmahan |last5=Zalloua |first5=Pierre |last6=Comas |first6=David |date=2019-11-18 |title=Heterogeneity in Palaeolithic Population Continuity and Neolithic Expansion in North Africa |url=https://www.sciencedirect.com/science/article/pii/S0960982219312412 |journal=Current Biology |language=en |volume=29 |issue=22 |pages=3953–3959.e4 |doi=10.1016/j.cub.2019.09.050 |issn=0960-9822}}</ref> A study (Pickrell et al. 2014) indicated that Western Eurasian ancestry eventually arrived through [[Northeast Africa]] (particularly the Horn of Africa) to [[Southern Africa]].<ref name=":3">{{cite journal |display-authors=6 |vauthors=Pickrell JK, Patterson N, Loh PR, Lipson M, Berger B, Stoneking M, Pakendorf B, Reich D |date=February 2014 |title=Ancient west Eurasian ancestry in southern and eastern Africa |journal=Proceedings of the National Academy of Sciences of the United States of America |volume=111 |issue=7 |pages=2632–2637 |arxiv=1307.8014 |bibcode=2014PNAS..111.2632P |doi=10.1073/pnas.1313787111 |pmc=3932865 |pmid=24550290 |doi-access=free}}</ref>
Another study (Ramsay ''et al.'' 2018) also found evidence for significant Western Eurasian admixture in various parts of Africa, from both ancient and more recent migrations, being highest among populations from [[North Africa|Northern Africa]], and some groups of the [[Horn of Africa]]:<ref name="Ramsay">{{cite journal |vauthors=Choudhury A, Aron S, Sengupta D, Hazelhurst S, Ramsay M |title=African genetic diversity provides novel insights into evolutionary history and local adaptations |journal=Human Molecular Genetics |volume=27 |issue=R2 |pages=R209–R218 |date=August 2018 |pmid=29741686 |pmc=6061870 |doi=10.1093/hmg/ddy161}}</ref>
<blockquote>
In addition to the intrinsic diversity within the continent due to population structure and isolation, migration of Eurasian populations into Africa has emerged as a critical contributor to the genetic diversity. These migrations involved the influx of different Eurasian populations at different times and to different parts of Africa. Comprehensive characterization of the details of these migrations through genetic studies on existing populations could help to explain the strong genetic differences between some geographically neighbouring populations.
This distinctive Eurasian admixture appears to have occurred over at least three time periods with ancient admixture in central west Africa (e.g. Yoruba from Nigeria) occurring between ~7.5 and 10.5 kya, older admixture in east Africa (e.g. Ethiopia) occurring between ~2.4 and 3.2 kya and more recent admixture between ~0.15 and 1.5 kya in some east African (e.g. Kenyan) populations.
Subsequent studies based on LD decay and haplotype sharing in an extensive set of African and Eurasian populations confirmed the presence of Eurasian signatures in west, east and southern Africans. In the west, in addition to Niger-Congo speakers from The Gambia and Mali, the Mossi from Burkina Faso showed the oldest Eurasian admixture event ~7 kya. In the east, these analyses inferred Eurasian admixture within the last 4000 years in Kenya.</blockquote>
[[File:Expansion of Afroasiatic.svg|thumb|Proposed migration and expansion routes of the [[Afroasiatic languages]] according to the indigenous African origin model.]]
There is no definitive agreement on when or where the [[Afroasiatic homeland|original homeland]] of the [[Afroasiatic languages|Afroasiatic language family]] existed. Some have suggested that they were spread by people with largely West-Eurasian ancestry during the [[Neolithic Revolution]], towards Northern Africa and the Horn of Africa, outgoing from the [[Middle East]], specifically from the [[Levant]]. This hypothesis does not account for the domestication of plants [[Endemism|endemic]] to the Horn of Africa such as [[teff]], [[Ensete ventricosum|ensete]], and [[Guizotia abyssinica|niger seed]], nor does it account for the lack of evidence of intrusive agricultural populations or the cultivation of [[wheat]], [[barley]], or [[sorghum]] in that region prior to 3000 B.C.<ref>{{Cite book |last1=Clark |first1=JD |title=From Hunters to Farmers: The Causes and Consequences of Food Production in Africa |last2=Brandt |first2=SA |publisher=University of California Press |year=1984 |isbn=978-0520045743 |pages=180 |language=English}}</ref><ref>{{cite journal |vauthors=Diamond J, Bellwood P |date=April 2003 |title=Farmers and their languages: the first expansions |journal=Science |volume=300 |issue=5619 |pages=597–603 |bibcode=2003Sci...300..597D |doi=10.1126/science.1078208 |pmid=12714734 |s2cid=13350469}}</ref> Others argue that the first speakers of Afroasiatic ([[Proto-Afroasiatic]]) were based in Northeast Africa because that region includes the majority of the diversity of the Afroasiatic language family and has very diverse groups in close geographic proximity, sometimes considered a telltale sign for a linguistic geographic origin.<ref>{{Cite book |last=Campbell |first=Lyle |title=Historical Linguistics, Fourth Edition |publisher=The MIT Press |year=2021 |isbn=978-0262542180 |pages=399–400 |language=English}}</ref> A subset of the Proto-Afroasiatic population would have migrated to the [[Levant]] during the late [[Paleolithic]], merging with local West-Eurasians and resulting in a population which would later give rise to [[Natufian culture]], associated with the early development of [[agriculture]] and early Afroasiatic languages, or specifically pre-[[Proto-Semitic language|proto-Semitic]].<ref>{{Cite book |last1=Jarvie |title=Transition to Modernity: Essays on Power, Wealth and Belief |last2=Hall |publisher=Cambridge University Press |year=2005 |isbn=9780521022279 |pages=27 |language=English}}</ref><ref>{{Cite book |last=Shirai |first=Noriyuki |url=http://worldcat.org/oclc/852516752 |title=The archaeology of the first farmer-herders in Egypt: new insights into the Fayum Epipalaeolithic and Neolithic |date=2010 |publisher=Leiden University Press |isbn=978-90-485-1269-0 |oclc=852516752}}</ref>{{Page needed|date=March 2022}}<ref name="Early back-to-Africa migration into">{{cite journal |vauthors=Hodgson JA, Mulligan CJ, Al-Meeri A, Raaum RL |date=June 2014 |title=Early back-to-Africa migration into the Horn of Africa |journal=PLOS Genetics |volume=10 |issue=6 |pages=e1004393 |doi=10.1371/journal.pgen.1004393 |pmc=4055572 |pmid=24921250}}</ref><ref>{{Cite book |last=Blench |first=Roger |title=Archaeology, Language, and the African Past |publisher=AltaMira Press |year=2006 |isbn=978-0759104662 |pages=150–163 |language=English}}</ref><ref name=":4">{{Cite journal |last=Ehret |first=Christopher |date=1979 |title=On the Antiquity of Agriculture in Ethiopia |url=https://www.jstor.org/stable/181512 |journal=The Journal of African History |volume=20 |issue=2 |pages=161–177 |doi=10.1017/S002185370001700X |jstor=181512 |s2cid=162986221 |via=JSTOR}}</ref><ref>{{Cite book |last=Nöth |first=Winfried |title=Origins of Semiosis: Sign Evolution in Nature and Culture |publisher=De Gruyter Mouton |year=2011 |isbn=9781134816231 |pages=293 |language=English}}</ref>
[[File:E of Y-DNA migrations.png|thumb|Proposed migration routes of paternal lineage E.]]
While many studies conducted on Horn of Africa populations estimate a West-Eurasian admixture event around 3,000 years ago,<ref name=":0" /><ref name="Ramsay" /><ref name=":3" /><ref>{{Cite journal |last1=Molinaro |first1=Ludovica |last2=Montinaro |first2=Francesco |last3=Yelmen |first3=Burak |last4=Marnetto |first4=Davide |last5=Behar |first5=Doron M. |last6=Kivisild |first6=Toomas |last7=Pagani |first7=Luca |date=2019-12-11 |title=West Asian sources of the Eurasian component in Ethiopians: a reassessment |journal=Scientific Reports |language=en |volume=9 |issue=1 |pages=18811 |doi=10.1038/s41598-019-55344-y |pmid=31827175 |pmc=6906521 |bibcode=2019NatSR...918811M |issn=2045-2322}}</ref> a 2014 genome study by Hodgson et al. found a distinct West-Eurasian ancestral component among studied Afroasiatic-speaking groups in the Horn of Africa (and to a lesser extent in North Africa and [[Western Asia|West Asia]]), most prevalent among the [[Somalis|Somali]]. This ancestral component — dubbed "Ethio-Somali" — would have diverged from other non-African ancestries around 23,000 years ago and migrated back to Africa prior to developing agriculture, merging with the local indigenous lineages of the Horn of Africa. The authors propose that "Ethio-Somali" may have been a substantial ancestral component of the Proto-Afroasiatic-speaking population. A subsequent [[Mitochondrial DNA|mtDNA]] (Gandini et al. 2016) analysis has produced additional evidence in support of a pre-agricultural back-migration from West-Eurasia into the [[Horn of Africa]] with an estimated date of arrival into the Horn of Africa in the early [[Holocene]], possibly as a result of [[obsidian]] exchange networks across the Red Sea.<ref>{{Cite journal |last1=Gandini |first1=Francesca |last2=Achilli |first2=Alessandro |last3=Pala |first3=Maria |last4=Bodner |first4=Martin |last5=Brandini |first5=Stefania |last6=Huber |first6=Gabriela |last7=Egyed |first7=Balazs |last8=Ferretti |first8=Luca |last9=Gómez-Carballa |first9=Alberto |last10=Salas |first10=Antonio |last11=Scozzari |first11=Rosaria |date=2016-05-05 |title=Mapping human dispersals into the Horn of Africa from Arabian Ice Age refugia using mitogenomes |journal=Scientific Reports |language=en |volume=6 |issue=1 |pages=25472 |doi=10.1038/srep25472 |pmid=27146119 |pmc=4857117 |bibcode=2016NatSR...625472G |issn=2045-2322}}</ref> Hodgson et al. also confirmed the existence of an ancestral component indigenous to the Horn of Africa – "Ethiopic" (Hodgson et al.) or "Omotic" (Pagani et al.) – which is most prevalent among speakers of the [[Omotic languages|Omotic]] branch of Afroasiatic in southwestern Ethiopia.<ref name="Early back-to-Africa migration into" /><ref name=":0" /> This lineage is associated with that of a 4,500 year-old fossil (Mota) found in a cave in southwestern Ethiopia, which has high genetic affinity to modern [[Ethiopians|Ethiopian groups]], especially the [[Endogamy|endogamous]] blacksmith [[caste]] of the Omotic [[Ari people|Aari people]]. Like Mota, Aari blacksmiths do not show evidence for admixture with West-Eurasians, demonstrating a degree of population continuity in this region for at least 4,500 years. In a comparative analysis of Mota’s genome referencing modern populations, Gallego et al. (2016) concluded that the divergence of Omotic from other Afroasiatic languages may have resulted from the relative isolation of its speakers from external groups.<ref>{{Cite journal |date=2016-02-19 |title=Erratum for the Report "Ancient Ethiopian genome reveals extensive Eurasian admixture in Eastern Africa" (previously titled "Ancient Ethiopian genome reveals extensive Eurasian admixture throughout the African continent") by M. Gallego Llorente, E. R. Jones, A. Eriksson, V. Siska, K. W. Arthur, J. W. Arthur, M. C. Curtis, J. T. Stock, M. Coltorti, P. Pieruccini, S. Stretton, F. Brock, T. Higham, Y. Park, M. Hofreiter, D. G. Bradley, J. Bhak, R. Pinhasi, A. Manica |url=https://pubmed.ncbi.nlm.nih.gov/26912899/ |journal=Science |volume=351 |issue=6275 |pages=aaf3945 |doi=10.1126/science.aaf3945 |issn=1095-9203 |pmid=26912899}}</ref> In an analysis of 68 Ethiopian ethnic groups, also referencing Mota, Lopez et al. (2021) revealed that groups belonging to the Omotic, Cushitic, and Semitic branches of Afro-Asiatic show high genetic similarity to each other on average. The data also support widespread recent intermixing among ethnic groups.<ref>{{Cite journal |last1=López |first1=Saioa |last2=Tarekegn |first2=Ayele |last3=Band |first3=Gavin |last4=van Dorp |first4=Lucy |last5=Bird |first5=Nancy |last6=Morris |first6=Sam |last7=Oljira |first7=Tamiru |last8=Mekonnen |first8=Ephrem |last9=Bekele |first9=Endashaw |last10=Blench |first10=Roger |last11=Thomas |first11=Mark G. |date=2021-06-11 |title=Evidence of the interplay of genetics and culture in Ethiopia |journal=Nature Communications |language=en |volume=12 |issue=1 |pages=3581 |doi=10.1038/s41467-021-23712-w |pmid=34117245 |pmc=8196081 |bibcode=2021NatCo..12.3581L |issn=2041-1723}}</ref> Linguist [[Roger Blench]] proposed southwestern Ethiopia as the most likely homeland of Afroasiatic, due in part to the high internal diversification of the Omotic branch spoken in that region.<ref name="BlenchAfricanPast" /> In addition, [[Human Y-chromosome DNA haplogroup|Y-haplogroup]] sub-lineage [[Haplogroup E-M215 (Y-DNA)|E-M215]] (also known as "E1b1b) and its derivative E-M35 are quite common among Afroasiatic speakers and southwestern Ethiopia is a plausible source of these haplogroups.<ref name=":1">{{Cite journal |last=Shriner |first=Daniel |date=2018 |title=Re-analysis of Whole Genome Sequence Data From 279 Ancient Eurasians Reveals Substantial Ancestral Heterogeneity |journal=Frontiers in Genetics |volume=9 |page=268 |doi=10.3389/fgene.2018.00268 |issn=1664-8021 |pmc=6062619 |pmid=30079081 |quote=and a sub-Saharan African component in Natufians that localizes to present-day southern Ethiopia. |doi-access=free}}</ref> The linguistic group and carriers of this lineage have a high probability to have arisen and dispersed together from Northeast Africa in the [[Mesolithic]], plausibly having already developed [[subsistence pattern]]s of [[pastoralism]] and intensive plant usage and collection.<ref>{{Cite journal |last1=Underhill |first1=P. A. |last2=Passarino |first2=G. |last3=Lin |first3=A. A. |last4=Shen |first4=P. |last5=Mirazón Lahr |first5=M. |last6=Foley |first6=R. A. |last7=Oefner |first7=P. J. |last8=Cavalli-Sforza |first8=L. L. |date=January 2001 |title=The phylogeography of Y chromosome binary haplotypes and the origins of modern human populations |journal=Annals of Human Genetics |volume=65 |issue=Pt 1 |pages=43–62 |doi=10.1046/j.1469-1809.2001.6510043.x |issn=0003-4800 |pmid=11415522 |s2cid=9441236}}</ref><ref>{{Cite journal |last=Ibrahim |first=Muntaser E. |date=2021-04-26 |title=Genetic diversity of the Sudanese: insights on origin and implications for health |journal=Human Molecular Genetics |volume=30 |issue=R1 |pages=R37–R41 |doi=10.1093/hmg/ddab028 |issn=1460-2083 |pmc=8223596 |pmid=33864377}}</ref><ref>{{Cite journal |last1=Ehret |first1=Christopher |last2=Keita |first2=S. O. Y. |last3=Newman |first3=Paul |date=2004-12-03 |title=The origins of Afroasiatic |journal=Science |volume=306 |issue=5702 |pages=1680; author reply 1680 |doi=10.1126/science.306.5702.1680c |issn=1095-9203 |pmid=15576591 |s2cid=8057990}}</ref><ref>{{Cite journal |last=Ehret |first=Christopher |date=1979 |title=On the Antiquity of Agriculture in Ethiopia |journal=The Journal of African History |volume=20 |issue=2 |pages=161–177 |doi=10.1017/S002185370001700X |jstor=181512 |s2cid=162986221 |issn=0021-8537}}</ref> According to historian and linguist [[Christopher Ehret]], the form of intensive plant collection practiced by the Proto-Afroasiatic population in Northeast Africa may have been a precursor to the agricultural practices that would later independently develop in the [[Fertile Crescent]] and the Horn of Africa.<ref name=":4"/><ref>{{Citation |last1=Bultosa |first1=G. |title=TEFF |date=2004-01-01 |url=https://www.sciencedirect.com/science/article/pii/B0127654909001725 |encyclopedia=Encyclopedia of Grain Science |pages=281–290 |editor-last=Wrigley |editor-first=Colin |place=Oxford |publisher=Elsevier |language=en |doi=10.1016/b0-12-765490-9/00172-5 |isbn=978-0-12-765490-4 |access-date=2022-03-29 |last2=Taylor |first2=J. R. N.}}</ref><ref>{{Cite journal |last1=Schlebusch |first1=Carina M. |last2=Jakobsson |first2=Mattias |date=2018-08-31 |title=Tales of Human Migration, Admixture, and Selection in Africa |journal=Annual Review of Genomics and Human Genetics |volume=19 |pages=405–428 |doi=10.1146/annurev-genom-083117-021759 |issn=1545-293X |pmid=29727585 |s2cid=19155657}}</ref> A 2018 re-analysis of autosomal DNA using modern populations as a reference found that the ancient Natufian samples of the Levant harbored 6.8% Omotic-related ancestry.<ref name=":1"/> A 2015 study by Dobon et al. identified an autosomal ancestral component that is commonly found among modern Afroasiatic-speaking populations (as well as [[Nubians]]) in Northeast Africa. This component, which peaks among [[Copts]] in [[Sudan]] but is not found in [[Egyptians]] or [[Qataris]], appears alongside a component that defines [[Nilo-Saharan languages|Nilo-Saharan]] speakers of southwestern Sudan and [[South Sudan]].<ref>{{cite journal |display-authors=6 |vauthors=Dobon B, Hassan HY, Laayouni H, Luisi P, Ricaño-Ponce I, Zhernakova A, Wijmenga C, Tahir H, Comas D, Netea MG, Bertranpetit J |date=May 2015 |title=The genetics of East African populations: a Nilo-Saharan component in the African genetic landscape |journal=Scientific Reports |volume=5 |pages=9996 |bibcode=2015NatSR...5E9996D |doi=10.1038/srep09996 |pmc=4446898 |pmid=26017457}}</ref> A 2017 paper by Arauna et al. analyzing existing genetic data obtained from [[North Africa|Northern African]] populations, such as [[Berbers]], described them as a mosaic of Middle Eastern, European, and [[Sub-Saharan Africa]]n-related ancestries.<ref>{{Citation |vauthors=Arauna LR, Comas D |title=Genetic Heterogeneity between Berbers and Arabs |date=2017 |work=eLS |pages=1–7 |publisher=John Wiley & Sons, Ltd |language=en |doi=10.1002/9780470015902.a0027485 |isbn=978-0-470-01590-2}}</ref>
[[File:Austronesia with hypothetical greatest expansion extent (Blench, 2009) 01.png|thumb|Austronesian expansion, outgoing from [[Taiwan]] and the northern [[Philippines]].]]
Specific East Asian-related ancestry is found among the [[Malagasy languages|Malagasy speakers]] of [[Madagascar]] at a medium frequency. The presence of this East Asian-related ancestry is mostly linked to the [[Austronesian peoples]] expansion from [[Southeast Asia]].<ref>{{Cite journal |vauthors=Dewar RE, Wright HT |date=1993-12-01|title=The culture history of Madagascar |journal=Journal of World Prehistory |language=en |volume=7 |issue=4 |pages=417–466 |doi=10.1007/BF00997802 |hdl=2027.42/45256 |s2cid=21753825 |hdl-access=free}}</ref><ref>{{cite journal | vauthors = Burney DA, Burney LP, Godfrey LR, Jungers WL, Goodman SM, Wright HT, Jull AJ | title = A chronology for late prehistoric Madagascar | journal = Journal of Human Evolution | volume = 47 | issue = 1–2 | pages = 25–63 | date = 2004-07-01 | pmid = 15288523 | doi = 10.1016/j.jhevol.2004.05.005 }}</ref><ref>{{cite journal | vauthors = Pierron D, Razafindrazaka H, Pagani L, Ricaut FX, Antao T, Capredon M, Sambo C, Radimilahy C, Rakotoarisoa JA, Blench RM, Letellier T, Kivisild T | display-authors = 6 | title = Genome-wide evidence of Austronesian-Bantu admixture and cultural reversion in a hunter-gatherer group of Madagascar | journal = Proceedings of the National Academy of Sciences of the United States of America | volume = 111 | issue = 3 | pages = 936–941 | date = January 2014 | pmid = 24395773 | pmc = 3903192 | doi = 10.1073/pnas.1321860111 | doi-access = free | bibcode = 2014PNAS..111..936P }}</ref><ref name="Kusuma_2016">{{cite journal | vauthors = Kusuma P, Brucato N, Cox MP, Pierron D, Razafindrazaka H, Adelaar A, Sudoyo H, Letellier T, Ricaut FX | display-authors = 6 | title = Contrasting Linguistic and Genetic Origins of the Asian Source Populations of Malagasy | journal = Scientific Reports | volume = 6 | issue = 1 | pages = 26066 | date = May 2016 | pmid = 27188237 | pmc = 4870696 | doi = 10.1038/srep26066 | bibcode = 2016NatSR...626066K }}</ref> The peoples of [[Borneo]] were identified to resemble the East Asian voyagers, who arrived on Madagascar. East Asian ancestry among Malagasy people was estimated at a mean average of 33%, but as high as ~75% among some Highlander groups and upper caste groups.<ref name="Heiske"/><ref>{{Cite web |last=Nicolas |first=Brucato |display-authors=etal |date=4 February 2019 |title=Evidence of Austronesian Genetic Lineages in East Africa and South Arabia: Complex Dispersal from Madagascar and Southeast Asia |url=https://academic.oup.com/gbe/article/11/3/748/5306180 |website=Genome Biology and Evolution, Volume 11, Issue 3}}</ref><ref name="Kusuma_2016" />
A 2020 study by Chen et al. analyzed 2,504 African samples from all over Africa, and found archaic Neanderthal ancestry, among all tested African samples at low frequency. They also identified a European-related (West-Eurasian) ancestry segment, which seems to largely correspond with the detected Neanderthal ancestry components. European-related admixture among Africans was estimated to be between ~0% to up to ~30%, with a peak among Northern Africans.<ref name="Identifying and Interpreting Appare"/>
According to the authors:<ref name="Identifying and Interpreting Appare"/>
{{Blockquote|text=These data are consistent with the hypothesis that back-migration contributed to the signal of Neanderthal ancestry in Africans. Furthermore, the data indicates that this back-migration came after the split of Europeans and East Asians, from a population related to the European lineage."}}
A 2021 paper by Hollfelder et al., concluded that West African [[Yoruba people]], which were previously used as "unadmixed reference population" for indigenous Africans, harbor minor levels of Neanderthal ancestry, which can be largely associated with back-migration of an "Ancestral European-like" source population.<ref name="The deep population history in Afri"/>
[[File:Map_of_African_admixture_in_European_populations.png|thumb|Map of [[Sub-Saharan African]] and specific [[North African]] admixture in European populations]]
Multiple studies found also evidence for geneflow of indigenous African ancestry towards Eurasia, specifically Europe and the Middle East. The analysis of 40 different West-Eurasian populations found African admixture at a frequency of 0% to up to ~15%.<ref>{{Cite journal |last1=Moorjani |first1=Priya |last2=Patterson |first2=Nick |last3=Hirschhorn |first3=Joel N. |last4=Keinan |first4=Alon |last5=Hao |first5=Li |last6=Atzmon |first6=Gil |last7=Burns |first7=Edward |last8=Ostrer |first8=Harry |last9=Price |first9=Alkes L. |last10=Reich |first10=David |date=April 2011 |title=The history of African gene flow into Southern Europeans, Levantines, and Jews |journal=PLOS Genetics |volume=7 |issue=4 |pages=e1001373 |doi=10.1371/journal.pgen.1001373 |issn=1553-7404 |pmc=3080861 |pmid=21533020}}</ref><ref>{{Cite journal |last1=Botigué |first1=Laura R. |last2=Henn |first2=Brenna M. |last3=Gravel |first3=Simon |last4=Maples |first4=Brian K. |last5=Gignoux |first5=Christopher R. |last6=Corona |first6=Erik |last7=Atzmon |first7=Gil |last8=Burns |first8=Edward |last9=Ostrer |first9=Harry |last10=Flores |first10=Carlos |last11=Bertranpetit |first11=Jaume |date=2013-07-16 |title=Gene flow from North Africa contributes to differential human genetic diversity in southern Europe |journal=Proceedings of the National Academy of Sciences of the United States of America |volume=110 |issue=29 |pages=11791–11796 |doi=10.1073/pnas.1306223110 |issn=0027-8424 |pmc=3718088 |pmid=23733930 |bibcode=2013PNAS..11011791B |doi-access=free }}</ref><ref>{{Cite journal |last1=Auton |first1=Adam |last2=Bryc |first2=Katarzyna |last3=Boyko |first3=Adam R. |last4=Lohmueller |first4=Kirk E. |last5=Novembre |first5=John |last6=Reynolds |first6=Andy |last7=Indap |first7=Amit |last8=Wright |first8=Mark H. |last9=Degenhardt |first9=Jeremiah D. |last10=Gutenkunst |first10=Ryan N. |last11=King |first11=Karen S. |date=May 2009 |title=Global distribution of genomic diversity underscores rich complex history of continental human populations |journal=Genome Research |volume=19 |issue=5 |pages=795–803 |doi=10.1101/gr.088898.108 |issn=1088-9051 |pmc=2675968 |pmid=19218534}}</ref><ref>{{Cite journal |last1=Reich |first1=David |last2=Price |first2=Alkes L. |last3=Patterson |first3=Nick |date=May 2008 |title=Principal component analysis of genetic data |url=https://www.nature.com/articles/ng0508-491 |journal=Nature Genetics |language=en |volume=40 |issue=5 |pages=491–492 |doi=10.1038/ng0508-491 |pmid=18443580 |s2cid=34837532 |issn=1546-1718}}</ref>
Evidence for minor geneflow from West or West-Central Africa into the [[Iberian Peninsula]] is supported by the presence of an African-specific mitochondrial haplogroup among one of four 4,000 year old samples.<ref>{{Cite journal |last1=González-Fortes |first1=G. |last2=Tassi |first2=F. |last3=Trucchi |first3=E. |last4=Henneberger |first4=K. |last5=Paijmans |first5=J. L. A. |last6=Díez-del-Molino |first6=D. |last7=Schroeder |first7=H. |last8=Susca |first8=R. R. |last9=Barroso-Ruíz |first9=C. |last10=Bermudez |first10=F. J. |last11=Barroso-Medina |first11=C. |last12=Bettencourt |first12=A. M. S. |last13=Sampaio |first13=H. A. |last14=Grandal-d'Anglade |first14=A. |last15=Salas |first15=A. |date=2019-01-30 |title=A western route of prehistoric human migration from Africa into the Iberian Peninsula |journal=Proceedings of the Royal Society B: Biological Sciences |volume=286 |issue=1895 |pages=20182288 |doi=10.1098/rspb.2018.2288 |issn=0962-8452 |pmc=6364581 |pmid=30963949}}</ref>
==Regional genomic overview==
===North Africa===
{{Main|Genetic history of North Africa}}
{{Further|DNA history of Egypt|Genetic studies on Moroccans}}
====Archaic Human DNA====
{{Further|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.<ref name="Bergström">{{cite journal | vauthors = Bergström A, Stringer C, Hajdinjak M, Scerri EM, Skoglund P | title = Origins of modern human ancestry | journal = Nature | volume = 590 | issue = 7845 | pages = 229–237 | date = February 2021 | pmid = 33568824 | doi = 10.1038/s41586-021-03244-5 | s2cid = 231883210 | bibcode = 2021Natur.590..229B }}</ref>
====Ancient DNA====
An early medieval sample from the 5th to 6th century CE found in northern France (Angers) was found to be similar to present-day North Africans.<ref>{{Cite journal |last=Alves |first=Isabel |last2=Giemza |first2=Joanna |last3=Blum |first3=Michael |last4=Bernhardsson |first4=Carolina |last5=Chatel |first5=Stéphanie |last6=Karakachoff |first6=Matilde |last7=Pierre |first7=Aude Saint |last8=Herzig |first8=Anthony F. |last9=Olaso |first9=Robert |last10=Monteil |first10=Martial |last11=Gallien |first11=Véronique |last12=Cabot |first12=Elodie |last13=Svensson |first13=Emma |last14=Bacq-Daian |first14=Delphine |last15=Baron |first15=Estelle |date=2022-02-04 |title=Genetic population structure across Brittany and the downstream Loire basin provides new insights on the demographic history of Western Europe |url=https://www.biorxiv.org/content/10.1101/2022.02.03.478491v1 |language=en |pages=2022.02.03.478491 |doi=10.1101/2022.02.03.478491v1.full}}</ref>
=====Egypt=====
[[Tomb of Two Brothers#The tomb owners|Khnum-aa]], [[Tomb of Two Brothers|Khnum-Nakht]], [[Tomb of Two Brothers#Coffins of Nakht-Ankh|Nakht-Ankh]] and JK2911 carried maternal [[Haplogroup M (mtDNA)#Haplogroup M1|haplogroup M1a1]].<ref name="Gad" /><ref name=":0">{{Cite journal |last1=Schuenemann |first1=Verena J. |last2=Peltzer |first2=Alexander |last3=Welte |first3=Beatrix |last4=van Pelt |first4=W. Paul |last5=Molak |first5=Martyna |last6=Wang |first6=Chuan-Chao |last7=Furtwängler |first7=Anja |last8=Urban |first8=Christian |last9=Reiter |first9=Ella |last10=Nieselt |first10=Kay |last11=Teßmann |first11=Barbara |date=2017-05-30 |title=Ancient Egyptian mummy genomes suggest an increase of Sub-Saharan African ancestry in post-Roman periods |journal=Nature Communications |volume=8 |issue=1 |page=15694 |doi=10.1038/ncomms15694 |pmid=28556824 |issn=2041-1723 |doi-access=free |bibcode=2017NatCo...815694S |pmc=5459999}}</ref>
[[Djehutynakht (10A)]] carried maternal haplogroup [[Haplogroup U (mtDNA)#Haplogroup U5|U5b2b5]].<ref name="Loreille 135">{{Cite journal |last1=Loreille |first1=Odile |last2=Ratnayake |first2=Shashikala |last3=Bazinet |first3=Adam L. |last4=Stockwell |first4=Timothy B. |last5=Sommer |first5=Daniel D. |last6=Rohland |first6=Nadin |last7=Mallick |first7=Swapan |last8=Johnson |first8=Philip L. F. |last9=Skoglund |first9=Pontus |last10=Onorato |first10=Anthony J. |last11=Bergman |first11=Nicholas H. |date=March 2018 |title=Biological Sexing of a 4000-Year-Old Egyptian Mummy Head to Assess the Potential of Nuclear DNA Recovery from the Most Damaged and Limited Forensic Specimens |journal=Genes |volume=9 |issue=3 |pages=135 |doi=10.3390/genes9030135 |pmid=29494531 |pmc=5867856 |doi-access=free}}</ref> JK2888 carried maternal haplogroup [[Haplogroup U (mtDNA)|U6a2]].<ref name=":0" />
[[Thuya]], [[Tiye]], Tutankhamen's mother, and Tutankhamen carried the maternal [[Haplogroup K (mtDNA)|haplogroup K]].<ref name="Gad" />
JK2134 carried maternal haplogroup [[Haplogroup J (mtDNA)|J1d]]<ref name=":0" /> and JK2887 carried maternal haplogroup [[Haplogroup J (mtDNA)|J2a1a1]].<ref name=":0" />
[[Amenhotep III]], [[Akhenaten]], and [[Tutankhamen]] carried the paternal [[haplogroup R1b]].<ref name="Gad">{{cite journal |vauthors=Gad YZ, Hassan NA, Mousa DM, Fouad FA, El-Sayed SG, Abdelazeem MA, Mahdy SM, Othman HY, Ibrahim DW, Khairat R, Ismail S |display-authors=6 |title=Insights from ancient DNA analysis of Egyptian human mummies: clues to disease and kinship |journal=Human Molecular Genetics |volume=30 |issue=R1 |pages=R24–R28 |date=April 2021 |pmid=33059357 |doi=10.1093/hmg/ddaa223 |doi-access=free}}</ref>
[[Ramesses III]] and "Unknown Man E", possibly [[Pentawere]], carried paternal [[haplogroup E1b1a]].<ref name="Gad" /><ref name="Hawass">{{cite journal |vauthors=Hawass Z, Ismail S, Selim A, Saleem SN, Fathalla D, Wasef S, Gad AZ, Saad R, Fares S, Amer H, Gostner P, Gad YZ, Pusch CM, Zink AR |display-authors=6 |title=Revisiting the harem conspiracy and death of Ramesses III: anthropological, forensic, radiological, and genetic study |journal=BMJ |volume=345 |pages=e8268 |date=December 2012 |pmid=23247979 |doi=10.1136/bmj.e8268 |hdl-access=free |s2cid=206896841 |hdl=10072/62081}}</ref><ref name="Gourdine">{{cite journal | vauthors = Gourdine JP, Keita S, Gourdine JL, Anselin A |title=Ancient Egyptian Genomes from northern Egypt: Further discussion |url=https://www.researchgate.net/publication/327065612 |journal=Nature Communications}}</ref>
JK2134 and JK2911 carried paternal haplogroup [[Haplogroup J (Y-DNA)|J]].<ref name=":0" />
[[Takabuti]] carried maternal haplogroup [[Haplogroup H (mtDNA)|H4a1]]<ref>{{Cite web |url=https://www.manchester.ac.uk/discover/news/shocking-truth-behind-takabutis-death-revealed/ |title=Shocking truth behind Takabuti's death revealed |language=en |access-date=2020-01-29}}</ref> and YM:KMM A 63 carried maternal haplogroup [[Haplogroup HV. (mtDNA)|HV]]<ref name=":10">{{Cite journal |last1=Oras |first1=Ester |last2=Anderson |first2=Jaanika |last3=Tõrv |first3=Mari |last4=Vahur |first4=Signe |last5=Rammo |first5=Riina |last6=Remmer |first6=Sünne |last7=Mölder |first7=Maarja |last8=Malve |first8=Martin |last9=Saag |first9=Lehti |last10=Saage |first10=Ragnar |last11=Teearu-Ojakäär |first11=Anu |date=2020-01-16 |title=Multidisciplinary investigation of two Egyptian child mummies curated at the University of Tartu Art Museum, Estonia (Late/Graeco-Roman Periods) |journal=PLOS ONE |language=en |volume=15 |issue=1 |pages=e0227446 |doi=10.1371/journal.pone.0227446 |pmid=31945091 |pmc=6964855 |bibcode=2020PLoSO..1527446O |issn=1932-6203 |doi-access=free}}</ref>
OM:KMM A 64 carried maternal haplogroup [[Haplogroup T (mtDNA)|T2c1a]].<ref name=":10" />
JK2888 carried paternal haplogroup [[Haplogroup E-M215 (Y-DNA)|E1b1b1a1b2]].<ref name=":0" />
=====Libya=====
At Takarkori rockshelter, in [[Libya]], two naturally [[mummified]] women, dated to the [[Pastoral Period#Middle Pastoral Period 2|Middle Pastoral Period]] (7000 BP), carried [[Basal (phylogenetics)|basal]] maternal [[Haplogroup N (mtDNA)|haplogroup N]].<ref name="Vai">{{cite journal | vauthors = Vai S, Sarno S, Lari M, Luiselli D, Manzi G, Gallinaro M, Mataich S, Hübner A, Modi A, Pilli E, Tafuri MA, Caramelli D, di Lernia S | display-authors = 6 | title = Ancestral mitochondrial N lineage from the Neolithic 'green' Sahara | journal = Scientific Reports | volume = 9 | issue = 1 | pages = 3530 | date = March 2019 | pmid = 30837540 | pmc = 6401177 | doi = 10.1038/s41598-019-39802-1 | bibcode = 2019NatSR...9.3530V }}</ref>
=====Morocco=====
A 2018 study by Van de Loorsdrecht et al. found that of seven samples of [[Taforalts]] of [[Morocco]], [[radiocarbon dated]] to between 15,100 cal BP and 13,900 cal BP, six were found to carry maternal haplogroup [[Haplogroup U (mtDNA)|U6a]], and one was found to carry maternal haplogroup [[Haplogroup M (mtDNA)|M1b]]. Six of six males were found to carry paternal haplogroup [[Haplogroup E-M215 (Y-DNA)|E1b1b]]. They were found to harbor 63.5% [[Natufian]]-related ancestry and 36.5% [[Sub-Saharan African]]-related ancestry. The Sub-Saharan component is most strongly drawn out by modern West African groups such as the [[Yoruba people|Yoruba]] and the [[Mende people|Mende]]. The samples also contain an additional affinity to South, Central, and East African outgroups that cannot be explained by any known ancient or modern populations.<ref name="Van De Loosdrecht">{{cite journal | vauthors = van de Loosdrecht M, Bouzouggar A, Humphrey L, Posth C, Barton N, Aximu-Petri A, Nickel B, Nagel S, Talbi EH, El Hajraoui MA, Amzazi S, Hublin JJ, Pääbo S, Schiffels S, Meyer M, Haak W, Jeong C, Krause J | display-authors = 6 | title = Pleistocene North African genomes link Near Eastern and sub-Saharan African human populations | journal = Science | volume = 360 | issue = 6388 | pages = 548–552 | date = May 2018 | pmid = 29545507 | doi = 10.1126/science.aar8380 | s2cid = 206666517 | doi-access = free | bibcode = 2018Sci...360..548V }}</ref> When projected onto a principal component analysis graph of African and west Eurasian populations, the Taforalt individuals form a distinct cluster in an intermediate position between present-day North Africans [e.g., [[Berbers|Amazighes]] (Berbers), Mozabites, and Saharawis] and East Africans (e.g., [[Afar people|Afars]], Oromos, and Somalis).<ref name="Van De Loosdrecht"/> Another study (Jeong 2020) comparing the Taforalt people of the [[Iberomaurusian]] culture to modern populations found that the Taforalt Sub-Saharan African genetic component may be best represented by modern West Africans (e.g. Yoruba).<ref name="Jeong" />
====Y-Chromosomal DNA====
{{Further|Y-DNA haplogroups in populations of North Africa}}
====Mitochondrial DNA====
Mitochondrial haplogroups L3, M, and N are found among [[Sudan#Ethnic groups|Sudanese peoples]] (e.g., [[Beja people|Beja]], [[Nilotic peoples|Nilotics]], [[Nuba]], [[Nubians]]), who have no known interaction (e.g., history of migration/admixture) with Europeans or Asians; rather than having developed in a post-Out-of-Africa migration context, mitochondrial macrohaplogroup L3/M/N and its subsequent development into distinct mitochondrial haplogroups (e.g., [[Haplogroup L3]], [[Haplogroup M (mtDNA)|Haplogroup M]], [[Haplogroup N (mtDNA)|Haplogroup N]]) may have occurred in [[East Africa]] at a time that considerably predates the Out-of-Africa migration event of 50,000 BP.<ref name="Osman">{{cite journal |vauthors=Osman MM, Hassan HY, Elnour MA, Makkan H, Gebremeskel EI, Gais T, Koko ME, Soodyall H, Ibrahim ME | display-authors = 6 |title=Mitochondrial HVRI and whole mitogenome sequence variations portray similar scenarios on the genetic structure and ancestry of northeast Africans |url=http://81.95.108.158/return-files/Mitochondrial%20HVRI.pdf |journal=Meta Gene}}</ref>
====Autosomal DNA====
====Medical DNA====
=====Lactase Persistence=====
[[Neolithic]] [[agriculturalists]], who may have resided in [[Northeast Africa]] and the [[Near East]], may have been the source population for [[lactase persistence]] variants, including –13910*T, and may have been subsequently supplanted by later migrations of peoples.<ref name="Priehodová">{{cite journal |vauthors=Priehodová E, Austerlitz F, Čížková M, Nováčková J, Ricaut FX, Hofmanová Z, Schlebusch CM, Černý V | display-authors = 6 | title = Sahelian pastoralism from the perspective of variants associated with lactase persistence | journal = American Journal of Physical Anthropology | volume = 173 | issue = 3 | pages = 423–436 | date = November 2020 | pmid = 32812238 | doi = 10.1002/ajpa.24116 | s2cid = 221179656 | url = https://hal.archives-ouvertes.fr/hal-02919786/file/ajpa_ms_final.pdf }}</ref> The [[Sub-Saharan]] [[West African]] Fulani, the [[North African]] [[Tuareg]], and [[Early European Farmers|European agriculturalists]], who are descendants of these Neolithic agriculturalists, share the lactase persistence variant –13910*T.<ref name="Priehodová" /> While shared by Fulani and Tuareg herders, compared to the Tuareg variant, the Fulani variant of –13910*T has undergone a longer period of haplotype differentiation.<ref name="Priehodová" /> The [[Fulani]] lactase persistence variant –13910*T may have spread, along with cattle [[pastoralism]], between 9686 BP and 7534 BP, possibly around 8500 BP; corroborating this timeframe for the Fulani, by at least 7500 BP, there is evidence of herders engaging in the act of [[milking]] in the Central [[Sahara]].<ref name="Priehodová" />
===West Africa===
====Archaic Human DNA====
{{Further|Interbreeding between archaic and modern humans#Archaic African hominins}}
Archaic traits found in human fossils of [[West Africa]] (e.g., [[Iwo Eleru skull|Iho Eleru fossils]], which dates to 13,000 BP) and [[Central Africa]] (e.g., [[Ishango]] fossils, which dates between 25,000 BP and 20,000 BP) may have developed as a result of admixture between archaic humans and modern humans or may be evidence of late-persisting [[Early modern human#Early Homo sapiens|early modern humans]].<ref name="Bergström"/> 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.<ref name="Bergström" />
====Ancient DNA====
As of 2017, human ancient DNA has not been found in the region of [[Western Africa]].<ref name="Scerri">{{cite web | vauthors = Scerri E |title= The Stone Age Archaeology of West Africa |url=https://ora.ox.ac.uk/objects/uuid:1442d1c1-7f76-454d-97db-8fd52a4c6c1e/download_file?file_format=pdf&safe_filename=The%2BStone%2BAge%2BArchaeology%2Bof%2BWest%2BAfrica.pdf&type_of_work=Book+section |website=Oxford University Research Archive |publisher=Oxford University Press}}</ref> As of 2020, human ancient DNA has not been forthcoming in the region of Western Africa.<ref name="Jeong">{{cite book | vauthors = Jeong C |chapter=Current Trends in Ancient DNA Study |chapter-url=https://link.springer.com/referenceworkentry/10.1007%2F978-981-15-1614-6_10-1 |year=2020 |pages=1–16 |title=The Handbook of Mummy Studies|publisher=Springer |doi=10.1007/978-981-15-1614-6_10-1 |isbn=978-981-15-1614-6 |s2cid=226555687 }}</ref>
In 4000 BP, there may have been a population that traversed from [[Africa]] (e.g., [[West Africa]] or West-[[Central Africa]]), through the [[Strait of Gibraltar]], into the [[Iberian peninsula]], where admixing between Africans and Iberians (e.g., of northern [[Portugal]], of southern [[Spain]]) occurred.<ref name="RSP">{{cite journal | vauthors = González-Fortes G, Tassi F, Trucchi E, Henneberger K, Paijmans JL, Díez-Del-Molino D, Schroeder H, Susca RR, Barroso-Ruíz C, Bermudez FJ, Barroso-Medina C, Bettencourt AM, Sampaio HA, Grandal-d'Anglade A, Salas A, de Lombera-Hermida A, Fabregas Valcarce R, Vaquero M, Alonso S, Lozano M, Rodríguez-Alvarez XP, Fernández-Rodríguez C, Manica A, Hofreiter M, Barbujani G | display-authors = 6 | title = A western route of prehistoric human migration from Africa into the Iberian Peninsula | journal = Proceedings. Biological Sciences | volume = 286 | issue = 1895 | pages = 20182288 | date = January 2019 | pmid = 30963949 | pmc = 6364581 | doi = 10.1098/rspb.2018.2288 }}</ref>
In [[Granada]], a Muslim ([[Moors|Moor]]) of the [[Cordoba Caliphate]],<ref name="Olalde">{{cite journal | vauthors = Olalde I, Mallick S, Patterson N, Rohland N, Villalba-Mouco V, Silva M, Dulias K, Edwards CJ, Gandini F, Pala M, Soares P, Ferrando-Bernal M, Adamski N, Broomandkhoshbacht N, Cheronet O, Culleton BJ, Fernandes D, Lawson AM, Mah M, Oppenheimer J, Stewardson K, Zhang Z, Jiménez Arenas JM, Toro Moyano IJ, Salazar-García DC, Castanyer P, Santos M, Tremoleda J, Lozano M, García Borja P, Fernández-Eraso J, Mujika-Alustiza JA, Barroso C, Bermúdez FJ, Viguera Mínguez E, Burch J, Coromina N, Vivó D, Cebrià A, Fullola JM, García-Puchol O, Morales JI, Oms FX, Majó T, Vergès JM, Díaz-Carvajal A, Ollich-Castanyer I, López-Cachero FJ, Silva AM, Alonso-Fernández C, Delibes de Castro G, Jiménez Echevarría J, Moreno-Márquez A, Pascual Berlanga G, Ramos-García P, Ramos-Muñoz J, Vijande Vila E, Aguilella Arzo G, Esparza Arroyo Á, Lillios KT, Mack J, Velasco-Vázquez J, Waterman A, Benítez de Lugo Enrich L, Benito Sánchez M, Agustí B, Codina F, de Prado G, Estalrrich A, Fernández Flores Á, Finlayson C, Finlayson G, Finlayson S, Giles-Guzmán F, Rosas A, Barciela González V, García Atiénzar G, Hernández Pérez MS, Llanos A, Carrión Marco Y, Collado Beneyto I, López-Serrano D, Sanz Tormo M, Valera AC, Blasco C, Liesau C, Ríos P, Daura J, de Pedro Michó MJ, Diez-Castillo AA, Flores Fernández R, Francès Farré J, Garrido-Pena R, Gonçalves VS, Guerra-Doce E, Herrero-Corral AM, Juan-Cabanilles J, López-Reyes D, McClure SB, Merino Pérez M, Oliver Foix A, Sanz Borràs M, Sousa AC, Vidal Encinas JM, Kennett DJ, Richards MB, Werner Alt K, Haak W, Pinhasi R, Lalueza-Fox C, Reich D | display-authors = 6 | title = The genomic history of the Iberian Peninsula over the past 8000 years | journal = Science | volume = 363 | issue = 6432 | pages = 1230–1234 | date = March 2019 | pmid = 30872528 | pmc = 6436108 | doi = 10.1126/science.aav4040 | bibcode = 2019Sci...363.1230O | hdl = 10261/207967 }}</ref> who was of haplogroups [[Haplogroup E-M2|E1b1a1]] and [[Haplogroup H (mtDNA)#H1|H1+16189]],<ref name="Olalde II">{{cite journal | vauthors = Olalde I |title=Ancient individuals from the Iberian Peninsula included in this study |url=https://science.sciencemag.org/highwire/filestream/724016/field_highwire_adjunct_files/2/aav4040_TablesS1-S5.xlsx |journal=Science}}</ref><ref name="Olalde III">{{cite journal | vauthors = Olalde I, Mallick S, Patterson N, Rohland N, Villalba-Mouco V, Silva M, Dulias K, Edwards CJ, Gandini F, Pala M, Soares P, Ferrando-Bernal M, Adamski N, Broomandkhoshbacht N, Cheronet O, Culleton BJ, Fernandes D, Lawson AM, Mah M, Oppenheimer J, Stewardson K, Zhang Z, Jiménez Arenas JM, Toro Moyano IJ, Salazar-García DC, Castanyer P, Santos M, Tremoleda J, Lozano M, García Borja P, Fernández-Eraso J, Mujika-Alustiza JA, Barroso C, Bermúdez FJ, Viguera Mínguez E, Burch J, Coromina N, Vivó D, Cebrià A, Fullola JM, García-Puchol O, Morales JI, Oms FX, Majó T, Vergès JM, Díaz-Carvajal A, Ollich-Castanyer I, López-Cachero FJ, Silva AM, Alonso-Fernández C, Delibes de Castro G, Jiménez Echevarría J, Moreno-Márquez A, Pascual Berlanga G, Ramos-García P, Ramos-Muñoz J, Vijande Vila E, Aguilella Arzo G, Esparza Arroyo Á, Lillios KT, Mack J, Velasco-Vázquez J, Waterman A, Benítez de Lugo Enrich L, Benito Sánchez M, Agustí B, Codina F, de Prado G, Estalrrich A, Fernández Flores Á, Finlayson C, Finlayson G, Finlayson S, Giles-Guzmán F, Rosas A, Barciela González V, García Atiénzar G, Hernández Pérez MS, Llanos A, Carrión Marco Y, Collado Beneyto I, López-Serrano D, Sanz Tormo M, Valera AC, Blasco C, Liesau C, Ríos P, Daura J, de Pedro Michó MJ, Diez-Castillo AA, Flores Fernández R, Francès Farré J, Garrido-Pena R, Gonçalves VS, Guerra-Doce E, Herrero-Corral AM, Juan-Cabanilles J, López-Reyes D, McClure SB, Merino Pérez M, Oliver Foix A, Sanz Borràs M, Sousa AC, Vidal Encinas JM, Kennett DJ, Richards MB, Werner Alt K, Haak W, Pinhasi R, Lalueza-Fox C, Reich D | display-authors = 6 | title = The genomic history of the Iberian Peninsula over the past 8000 years | journal = Science | volume = 363 | issue = 6432 | pages = 1230–1234 | date = March 2019 | pmid = 30872528 | pmc = 6436108 | doi = 10.1126/science.aav4040 | bibcode = 2019Sci...363.1230O }}</ref> as well as estimated to date between 900 CE and 1000 CE, and a [[Morisco]],<ref name="Olalde" /> who was of [[Haplogroup L2 (mtDNA)#Haplogroup L2e|haplogroup L2e1]],<ref name="Olalde II" /><ref name="Olalde III" /> as well as estimated to date between 1500 CE and 1600 CE, were both found to be of Sub-Saharan West African (i.e., [[The Gambia#Ethnic groups|Gambian]]), [[Iberian Peninsula|Iberian]], and North African descent.<ref name="Olalde" />
====Y-Chromosomal DNA====
{{Further|Y-DNA haplogroups in populations of Sub-Saharan Africa}}
As a result of haplogroup D0, a basal branch of haplogroup DE, being found in three [[Nigerian]] men, it may be the case that [[haplogroup DE]], as well as its sublineages D0 and E, originated in [[Africa]].<ref name="Haber">{{cite journal | vauthors = Haber M, Jones AL, Connell BA, Arciero E, Yang H, Thomas MG, Xue Y, Tyler-Smith C | display-authors = 6 | title = A Rare Deep-Rooting D0 African Y-Chromosomal Haplogroup and Its Implications for the Expansion of Modern Humans Out of Africa | journal = Genetics | volume = 212 | issue = 4 | pages = 1421–1428 | date = August 2019 | pmid = 31196864 | pmc = 6707464 | doi = 10.1534/genetics.119.302368 }}</ref>
As of 19,000 years ago, Africans, bearing [[Haplogroup E-V38|haplogroup E1b1a-V38]], likely traversed across the [[Sahara]], from [[East Africa|east]] to [[West Africa|west]].<ref name="Shriner">{{cite journal | vauthors = Shriner D, Rotimi CN | title = Whole-Genome-Sequence-Based Haplotypes Reveal Single Origin of the Sickle Allele during the Holocene Wet Phase | journal = American Journal of Human Genetics | volume = 102 | issue = 4 | pages = 547–556 | date = April 2018 | pmid = 29526279 | pmc = 5985360 | doi = 10.1016/j.ajhg.2018.02.003 }}</ref> [[Haplogroup E-M2|E1b1a1-M2]] likely originated in [[West Africa]] or [[Central Africa]].<ref name="Trombetta">{{cite journal | vauthors = Trombetta B, D'Atanasio E, Massaia A, Ippoliti M, Coppa A, Candilio F, Coia V, Russo G, Dugoujon JM, Moral P, Akar N, Sellitto D, Valesini G, Novelletto A, Scozzari R, Cruciani F | display-authors = 6 | title = Phylogeographic Refinement and Large Scale Genotyping of Human Y Chromosome Haplogroup E Provide New Insights into the Dispersal of Early Pastoralists in the African Continent | journal = Genome Biology and Evolution | volume = 7 | issue = 7 | pages = 1940–1950 | date = June 2015 | pmid = 26108492 | pmc = 4524485 | doi = 10.1093/gbe/evv118 }}</ref>
====Mitochondrial DNA====
{{Further|Haplogroup L2 (mtDNA)}}
Around 18,000 BP, [[Mende people]], along with [[The Gambia#Ethnic groups|Gambian peoples]], grew in population size.<ref name="Miller">{{cite journal | vauthors = Miller EF, Manica A, Amos W | title = Global demographic history of human populations inferred from whole mitochondrial genomes | journal = Royal Society Open Science | volume = 5 | issue = 8 | pages = 180543 | date = August 2018 | pmid = 30225046 | pmc = 6124094 | doi = 10.1098/rsos.180543 | bibcode = 2018RSOS....580543M }}</ref>
In 15,000 BP, [[Niger-Congo]] speakers may have migrated from the [[Sahelian]] region of West Africa, along the [[Senegal River]], and introduced [[Haplogroup L2 (mtDNA)#Haplogroup L2a1|L2a1]] into [[North Africa]], resulting in modern [[Demographics of Mauritania#Ethnic groups|Mauritanian peoples]] and [[Berbers]] of [[Tunisia]] inheriting it.<ref name="Frigi">{{cite journal | vauthors = Frigi S, Cherni L, Fadhlaoui-Zid K, Benammar-Elgaaied A | title = Ancient local evolution of African mtDNA haplogroups in Tunisian Berber populations | journal = Human Biology | volume = 82 | issue = 4 | pages = 367–384 | date = August 2010 | pmid = 21082907 | doi = 10.3378/027.082.0402 | s2cid = 27594333 }}</ref>
Between 11,000 BP and 10,000 BP, [[Yoruba people]] and [[Esan people]] grew in population size.<ref name="Miller" />
Up to 11,000 years ago, Sub-Saharan West Africans, bearing [[Macro-haplogroup L (mtDNA)|macrohaplogroup L]] (e.g., [[Haplogroup L1 (mtDNA)#L1b|L1b1a11]], L1b1a6a, L1b1a8, L1b1a9a1, [[Haplogroup L2 (mtDNA)#Haplogroup L2a1|L2a1k]], [[Haplogroup L3 (mtDNA)#Subclade distribution|L3d1b1a]]), may have migrated through [[North Africa]] and into [[Europe]], mostly into [[southern Europe]] (e.g., [[Iberia]]).<ref name="Soares">{{cite journal | vauthors = Podgorná E, Soares P, Pereira L, Cerný V | title = The genetic impact of the lake chad basin population in North Africa as documented by mitochondrial diversity and internal variation of the L3e5 haplogroup | journal = Annals of Human Genetics | volume = 77 | issue = 6 | pages = 513–523 | date = November 2013 | pmid = 25069842 | doi = 10.1111/ahg.12040 | s2cid = 24672148 }}</ref>
====Autosomal DNA====
Between 2000 BP and 1500 BP, [[Nilo-Saharan]]-speakers may have migrated across the [[Sahel]], from [[East Africa]] into [[West Africa]], and admixed with [[Niger-Congo]]-speaking [[Berom people]].<ref name="Choudhury II">{{cite journal | vauthors = Choudhury A, Aron S, Botigué LR, Sengupta D, Botha G, Bensellak T, Wells G, Kumuthini J, Shriner D, Fakim YJ, Ghoorah AW, Dareng E, Odia T, Falola O, Adebiyi E, Hazelhurst S, Mazandu G, Nyangiri OA, Mbiyavanga M, Benkahla A, Kassim SK, Mulder N, Adebamowo SN, Chimusa ER, Muzny D, Metcalf G, Gibbs RA, Rotimi C, Ramsay M, Adeyemo AA, Lombard Z, Hanchard NA | display-authors = 6 | title = High-depth African genomes inform human migration and health | journal = Nature | volume = 586 | issue = 7831 | pages = 741–748 | date = October 2020 | pmid = 33116287 | doi = 10.1038/s41586-020-2859-7 | pmc = 7759466 | bibcode = 2020Natur.586..741C }}</ref>
A 2019 study on [[Fulani people]] by Fan et al., found that the Fulani show genetic affinity to isolated [[Afroasiatic languages|Afroasiatic-speaking]] groups in [[East Africa|Eastern Africa]], specifically [[Omotic languages|Omotic-speakers]] such as the [[Aari people]]. While the Fulani have nearly exclusive indigenous African ancestry (defined by West and East African ancestry), they also show traces of West-Eurasian-like admixture, supporting an ancestral homeland somewhere in North or Eastern Africa, and westwards expansion during the Neolithic, possibly caused by the arrival and expansion of West-Eurasian-related groups.<ref>{{Cite journal |last1=Fan |first1=Shaohua |last2=Kelly |first2=Derek E. |last3=Beltrame |first3=Marcia H. |last4=Hansen |first4=Matthew E. B. |last5=Mallick |first5=Swapan |last6=Ranciaro |first6=Alessia |last7=Hirbo |first7=Jibril |last8=Thompson |first8=Simon |last9=Beggs |first9=William |last10=Nyambo |first10=Thomas |last11=Omar |first11=Sabah A. |date=2019-04-26 |title=African evolutionary history inferred from whole genome sequence data of 44 indigenous African populations |url=https://doi.org/10.1186/s13059-019-1679-2 |journal=Genome Biology |volume=20 |issue=1 |pages=82 |doi=10.1186/s13059-019-1679-2 |issn=1474-760X |pmc=6485071 |pmid=31023338}}</ref>
====Medical DNA====
=====Sickle Cell=====
Amid the Green Sahara, the mutation for [[sickle cell]] originated in the [[Sahara]]<ref name="Shriner" /> or in the [[Northwestern Congolian lowland forests|northwest forest]] region of western Central Africa (e.g., Cameroon)<ref name="Shriner" /><ref name="Esoh">{{cite journal | vauthors = Esoh K, Wonkam A | title = Evolutionary history of sickle-cell mutation: implications for global genetic medicine | journal = Human Molecular Genetics | volume = 30 | issue = R1 | pages = R119–R128 | date = April 2021 | pmid = 33461216 | pmc = 8117455 | doi = 10.1093/hmg/ddab004 }}</ref> by at least 7,300 years ago,<ref name="Shriner" /><ref name="Esoh" /> though possibly as early as 22,000 years ago.<ref name="Laval">{{cite journal | vauthors = Laval G, Peyrégne S, Zidane N, Harmant C, Renaud F, Patin E, Prugnolle F, Quintana-Murci L | display-authors = 6 | title = Recent Adaptive Acquisition by African Rainforest Hunter-Gatherers of the Late Pleistocene Sickle-Cell Mutation Suggests Past Differences in Malaria Exposure | journal = American Journal of Human Genetics | volume = 104 | issue = 3 | pages = 553–561 | date = March 2019 | pmid = 30827499 | pmc = 6407493 | doi = 10.1016/j.ajhg.2019.02.007 }}</ref><ref name="Esoh" /> The ancestral sickle cell haplotype to modern haplotypes (e.g., [[Cameroon]]/[[Central African Republic]] and [[Benin]]/[[Senegal]] haplotypes) may have first arose in the ancestors of modern West Africans, bearing haplogroups [[Haplogroup E-M2#E1b1a1a1f|E1b1a1-L485]] and [[Haplogroup E-M2#E1b1a1a1g|E1b1a1-U175]] or their ancestral haplogroup E1b1a1-M4732.<ref name="Shriner" /> West Africans (e.g., [[Yoruba people|Yoruba]] and [[Esan people|Esan]] of Nigeria), bearing the Benin sickle cell haplotype, may have migrated through the [[Northeast Africa|northeastern region of Africa]] into the western region of [[Arabia]].<ref name="Shriner" /> West Africans (e.g., [[Mende people|Mende]] of Sierra Leone), bearing the Senegal sickle cell haplotype,<ref name="Yaseen">{{cite journal | vauthors = Yaseen NT, Al-Mamoori HS, Hassan MK |title=Sickle β-globin haplotypes among patients with sickle cell anemia in Basra, Iraq: A cross-sectional study | doi = 10.4103/ijh.ijh_20_19 |journal=Iraqi Journal of Hematology | date = January 2020 | volume = 9 | issue = 1 | pages = 23 |s2cid=216082225 }}</ref><ref name="Shriner" /> may have migrated into [[Mauritania]] (77% modern rate of occurrence) and Senegal (100%); they may also have migrated across the Sahara, into [[North Africa]], and from North Africa, into [[Southern Europe]], [[Turkey]], and a region near northern [[Iraq]] and southern Turkey.<ref name="Yaseen" /> Some may have migrated into and introduced the Senegal and Benin sickle cell haplotypes into [[Basra]], Iraq, where both occur equally.<ref name="Yaseen" /> West Africans, bearing the Benin sickle cell haplotype, may have migrated into the northern region of Iraq (69.5%), [[Jordan]] (80%), [[Lebanon]] (73%), [[Oman]] (52.1%), and [[Egypt]] (80.8%).<ref name="Yaseen" />
=====Schistosomes=====
According to Steverding (2020), while not definite: Near the [[African Great Lakes]], schistosomes (e.g., [[S. mansoni]], [[S. haematobium]]) underwent evolution.<ref name="Steverding">{{cite journal | vauthors = Steverding D | title = The spreading of parasites by human migratory activities | journal = Virulence | volume = 11 | issue = 1 | pages = 1177–1191 | date = December 2020 | pmid = 32862777 | pmc = 7549983 | doi = 10.1080/21505594.2020.1809963 }}</ref> Subsequently, there was an expansion alongside the Nile.<ref name="Steverding" /> From [[Egypt]], the presence of schistosomes may have expanded, via migratory [[Yoruba people]], into Western Africa.<ref name="Steverding" /> Thereafter, [[schistosomes]] may have expanded, via [[Bantu migration|migratory]] [[Bantu peoples]], into the rest of [[Sub-Saharan Africa]] (e.g., [[Southern Africa]], [[Central Africa]]).<ref name="Steverding" />
=====Thalassemia=====
Through pathways taken by [[Caravan (travellers)|caravan]]s, or via travel amid the [[Almoravid dynasty|Almovarid]] period, a population (e.g., [[Sub-Saharan]] [[West Africa#Demographics and languages|West Africans]]) may have introduced the –29 (A → G) [[Beta thalassemia|β-thalassemia]] mutation (found in notable amounts among [[African-Americans]]) into the [[North African]] region of [[Morocco]].<ref name="Agouti">{{cite journal | vauthors = Agouti I, Badens C, Abouyoub A, Levy N, Bennani M | title = Molecular basis of beta-thalassemia in Morocco: possible origins of the molecular heterogeneity | journal = Genetic Testing | volume = 12 | issue = 4 | pages = 563–568 | date = December 2008 | pmid = 18976160 | doi = 10.1089/gte.2008.0058 | s2cid = 46000591 }}</ref>
===Central Africa===
{{See also|African Pygmies#Genetics}}
====Archaic Human DNA====
{{Further|Interbreeding between archaic and modern humans#Archaic African hominins}}
Archaic traits found in human fossils of [[West Africa]] (e.g., [[Iwo Eleru skull|Iho Eleru fossils]], which dates to 13,000 BP) and [[Central Africa]] (e.g., [[Ishango]] fossils, which dates between 25,000 BP and 20,000 BP) may have developed as a result of admixture between archaic humans and modern humans or may be evidence of late-persisting [[Early modern human#Early Homo sapiens|early modern humans]].<ref name="Bergström"/> 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.<ref name="Bergström" />
====Ancient DNA====
In 4000 BP, there may have been a population that traversed from [[Africa]] (e.g., [[West Africa]] or West-[[Central Africa]]), through the [[Strait of Gibraltar]], into the [[Iberian peninsula]], where admixing between Africans and Iberians (e.g., of northern [[Portugal]], of southern [[Spain]]) occurred.<ref name="RSP"/>
=====Cameroon=====
[[West African hunter-gatherers]], in the region of western Central Africa (e.g., [[Shum Laka]], [[Cameroon]]), particularly between 8000 BP and 3000 BP, were found to be related to modern [[Central African foragers|Central African hunter-gatherers]] (e.g., [[Baka people (Cameroon and Gabon)|Baka]], [[Bakola]], [[Aka people|Biaka]], [[Bedzan people|Bedzan]]).<ref name="Lipson">{{cite journal | vauthors = Lipson M, Ribot I, Mallick S, Rohland N, Olalde I, Adamski N, Broomandkhoshbacht N, Lawson AM, López S, Oppenheimer J, Stewardson K, Asombang RN, Bocherens H, Bradman N, Culleton BJ, Cornelissen E, Crevecoeur I, de Maret P, Fomine FL, Lavachery P, Mindzie CM, Orban R, Sawchuk E, Semal P, Thomas MG, Van Neer W, Veeramah KR, Kennett DJ, Patterson N, Hellenthal G, Lalueza-Fox C, MacEachern S, Prendergast ME, Reich D | display-authors = 6 | title = Ancient West African foragers in the context of African population history | journal = Nature | volume = 577 | issue = 7792 | pages = 665–670 | date = January 2020 | pmid = 31969706 | pmc = 8386425 | doi = 10.1038/s41586-020-1929-1 | s2cid = 210862788 | bibcode = 2020Natur.577..665L }}</ref>
=====Democratic Republic of Congo=====
At Kindoki, in the [[Democratic Republic of Congo]], there were three individuals, dated to the [[protohistoric]] period (230 BP, 150 BP, 230 BP); one carried haplogroups [[Haplogroup E-M2#E1b1a1a1d|E1b1a1a1d1a2 (E-CTS99, E-CTS99)]] and [[Haplogroup L1 (mtDNA)#L1c|L1c3a1b]], another carried [[Haplogroup E-M96|haplogroup E (E-M96, E-PF1620)]], and the last carried haplogroups [[Haplogroup R1b#R1b (R-L278)|R1b1 (R-P25 1, R-M415)]] and [[Haplogroup L0 (mtDNA)#Distribution|L0a1b1a1]].<ref name="Wang">{{cite journal | vauthors = Wang K, Goldstein S, Bleasdale M, Clist B, Bostoen K, Bakwa-Lufu P, Buck LT, Crowther A, Dème A, McIntosh RJ, Mercader J, Ogola C, Power RC, Sawchuk E, Robertshaw P, Wilmsen EN, Petraglia M, Ndiema E, Manthi FK, Krause J, Roberts P, Boivin N, Schiffels S | display-authors = 6 | title = Ancient genomes reveal complex patterns of population movement, interaction, and replacement in sub-Saharan Africa | journal = Science Advances | volume = 6 | issue = 24 | pages = eaaz0183 | date = June 2020 | pmid = 32582847 | pmc = 7292641 | doi = 10.1126/sciadv.aaz0183 | bibcode = 2020SciA....6..183W }}</ref><ref name="Wang II">{{cite journal | vauthors = Wang K, Goldstein S, Bleasdale M, Clist B, Bostoen K, Bakwa-Lufu P, Buck LT, Crowther A, Dème A, McIntosh RJ, Mercader J, Ogola C, Power RC, Sawchuk E, Robertshaw P, Wilmsen EN, Petraglia M, Ndiema E, Manthi FK, Krause J, Roberts P, Boivin N, Schiffels S | display-authors = 6 | title = Ancient genomes reveal complex patterns of population movement, interaction, and replacement in sub-Saharan Africa | journal = Science Advances | volume = 6 | issue = 24 | pages = eaaz0183 | date = June 2020 | pmid = 32582847 | doi = 10.1126/sciadv.aaz0183 | pmc = 7292641 | bibcode = 2020SciA....6..183W }}</ref>
At Ngongo Mbata, in the [[Democratic Republic of Congo]], an individual, dated to the [[protohistoric]] period (220 BP), carried haplogroup [[Haplogroup L1 (mtDNA)#L1c|L1c3a]].<ref name="Wang" /><ref name="Wang II" />
At Matangai Turu Northwest, in the [[Democratic Republic of Congo]], an individual, dated to the [[Iron Age#Sub-Saharan Africa|Iron Age]] (750 BP), carried an undetermined haplogroup(s).<ref name="Wang" /><ref name="Wang II" />
====Y-Chromosomal DNA====
{{Further|Y-DNA haplogroups in populations of Sub-Saharan Africa}}
[[Haplogroup R1b#R1b1b (R-V88)|Haplogroup R-V88]] may have originated in western Central Africa (e.g., [[Equatorial Guinea]]), and, in the middle of the [[Holocene]], arrived in [[North Africa]] through population migration.<ref name="González-Santos">{{cite journal | vauthors = González M, Gomes V, López-Parra AM, Amorim A, Carracedo A, Sánchez-Diz P, Arroyo-Pardo E, Gusmão L | display-authors = 6 | title = The genetic landscape of Equatorial Guinea and the origin and migration routes of the Y chromosome haplogroup R-V88 | journal = European Journal of Human Genetics | volume = 21 | issue = 3 | pages = 324–331 | date = March 2013 | pmid = 22892526 | doi = 10.1038/ejhg.2012.167 | pmc = 3573200 }}</ref>
====Mitochondrial DNA====
{{Further|Haplogroup L1 (mtDNA)|Haplogroup L2 (mtDNA)}}
Mitochondrial [[Haplogroup L1 (mtDNA)|haplogroup L1c]] is strongly associated with pygmies, especially with [[Mbenga people|Bambenga]] groups.<ref name="Quintana08">{{cite journal | vauthors = Quintana-Murci L, Quach H, Harmant C, Luca F, Massonnet B, Patin E, Sica L, Mouguiama-Daouda P, Comas D, Tzur S, Balanovsky O, Kidd KK, Kidd JR, van der Veen L, Hombert JM, Gessain A, Verdu P, Froment A, Bahuchet S, Heyer E, Dausset J, Salas A, Behar DM | display-authors = 6 | title = Maternal traces of deep common ancestry and asymmetric gene flow between Pygmy hunter-gatherers and Bantu-speaking farmers | journal = Proceedings of the National Academy of Sciences of the United States of America | volume = 105 | issue = 5 | pages = 1596–1601 | date = February 2008 | pmid = 18216239 | pmc = 2234190 | doi = 10.1073/pnas.0711467105 | doi-access = free | bibcode = 2008PNAS..105.1596Q }}</ref> L1c prevalence was variously reported as: 100% in [[Gyele language|Ba-Kola]], 97% in [[Aka people|Aka (Ba-Benzélé)]], and 77% in [[Aka people|Biaka]],<ref name="Tishkoff">Sarah A. Tishkoff et al. 2007, [https://web.archive.org/web/20080517050807/http://mbe.oxfordjournals.org/cgi/content/full/24/10/2180 History of Click-Speaking Populations of Africa Inferred from mtDNA and Y Chromosome Genetic Variation.] Molecular Biology and Evolution 2007 24(10):2180-2195</ref> 100% of the [[Bedzan people|Bedzan (Tikar)]], 97% and 100% in the [[Baka (Cameroon and Gabon)|Baka people]] of [[Gabon]] and [[Cameroon]], respectively,<ref>Lluis Quintana-Murci et al. MtDNA diversity in Central Africa: from hunter-gathering to agriculturalism. CNRS-Institut Pasteur, Paris</ref> 97% in [[Bakoya]] (97%), and 82% in [[Bongo people (Gabon)|Ba-Bongo]].<ref name="Quintana08" /> Mitochondrial haplogroups [[Haplogroup L2 (mtDNA)|L2a]] and [[Haplogroup L0 (mtDNA)|L0a]] are prevalent among the [[Bambuti]].<ref name="Quintana08"/><ref>{{cite journal | vauthors = Silva M, Alshamali F, Silva P, Carrilho C, Mandlate F, Jesus Trovoada M, Černý V, Pereira L, Soares P | display-authors = 6 | title = 60,000 years of interactions between Central and Eastern Africa documented by major African mitochondrial haplogroup L2 | journal = Scientific Reports | volume = 5 | pages = 12526 | date = July 2015 | pmid = 26211407 | pmc = 4515592 | doi = 10.1038/srep12526 | bibcode = 2015NatSR...512526S }}</ref>
====Autosomal DNA====
Genetically, [[African pygmies]] have some key difference between them and [[Bantu peoples]].<ref>{{cite journal | vauthors = Jarvis JP, Scheinfeldt LB, Soi S, Lambert C, Omberg L, Ferwerda B, Froment A, Bodo JM, Beggs W, Hoffman G, Mezey J, Tishkoff SA | display-authors = 6 | title = Patterns of ancestry, signatures of natural selection, and genetic association with stature in Western African pygmies | journal = PLOS Genetics | volume = 8 | issue = 4 | pages = e1002641 | year = 2012 | pmid = 22570615 | pmc = 3343053 | doi = 10.1371/journal.pgen.1002641 }}</ref><ref>{{cite journal | vauthors = López Herráez D, Bauchet M, Tang K, Theunert C, Pugach I, Li J, Nandineni MR, Gross A, Scholz M, Stoneking M | display-authors = 6 | title = Genetic variation and recent positive selection in worldwide human populations: evidence from nearly 1 million SNPs | journal = PLOS ONE | volume = 4 | issue = 11 | pages = e7888 | date = November 2009 | pmid = 19924308 | pmc = 2775638 | doi = 10.1371/journal.pone.0007888 | doi-access = free | bibcode = 2009PLoSO...4.7888L }}</ref>
====Medical DNA====
Evidence suggests that, when compared to other [[Sub-Saharan]] African populations, [[African pygmy]] populations display unusually low levels of expression of the genes encoding for [[human growth hormone]] and [[Growth hormone receptor|its receptor]] associated with low [[Blood serum|serum]] levels of [[insulin-like growth factor-1]] and short stature.<ref name='Bozzola, 2009'>{{cite journal | vauthors = Bozzola M, Travaglino P, Marziliano N, Meazza C, Pagani S, Grasso M, Tauber M, Diegoli M, Pilotto A, Disabella E, Tarantino P, Brega A, Arbustini E | display-authors = 6 | title = The shortness of Pygmies is associated with severe under-expression of the growth hormone receptor | journal = Molecular Genetics and Metabolism | volume = 98 | issue = 3 | pages = 310–313 | date = November 2009 | pmid = 19541519 | doi = 10.1016/j.ymgme.2009.05.009 }}</ref>
===Eastern Africa===
====Archaic Human DNA====
{{Further|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.<ref name="Bergström"/>
====Ancient DNA====
=====Ethiopia=====
At [[Mota, Ethiopia|Mota]], in [[Ethiopia]], an individual, estimated to date to the 5th millennium BP, carried haplogroups [[Haplogroup E-P2|E1b1]] and [[Haplogroup L3 (mtDNA)#Subclade distribution|L3x2a]].<ref name="Llorente">{{cite journal | vauthors = Llorente MG, Jones ER, Eriksson A, Siska V, Arthur KW, Arthur JW, Curtis MC, Stock JT, Coltorti M, Pieruccini P, Stretton S, Brock F, Higham T, Park Y, Hofreiter M, Bradley DG, Bhak J, Pinhasi R, Manica A | display-authors = 6 | title = Ancient Ethiopian genome reveals extensive Eurasian admixture throughout the African continent | journal = Science | volume = 350 | issue = 6262 | pages = 820–822 | date = November 2015 | pmid = 26449472 | doi = 10.1126/science.aad2879 | bibcode = 2015Sci...350..820L | hdl = 2318/1661894 | s2cid = 25743789 }}</ref><ref name="Llorente II">{{cite journal | vauthors = Llorente MG, Jones ER, Eriksson A, Siska V, Arthur KW, Arthur JW, Curtis MC, Stock JT, Coltorti M, Pieruccini P, Stretton S, Brock F, Higham T, Park Y, Hofreiter M, Bradley DG, Bhak J, Pinhasi R, Manica A | display-authors = 6 |title=Supplementary Materials for Ancient Ethiopian genome reveals extensive Eurasian admixture in Eastern Africa |url=https://www.science.org/doi/10.1126/science.aad2879 |journal=Science | date = 13 November 2015 | volume = 350 | issue = 6262 | pages = 820–822 | doi = 10.1126/science.aad2879 | pmid = 26449472 | bibcode = 2015Sci...350..820L | hdl = 2318/1661894 | s2cid = 25743789 }}</ref> 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.<ref name="Hellenthal">{{cite journal | vauthors = Hellenthal G, Bird N, Morris S | title = Structure and ancestry patterns of Ethiopians in genome-wide autosomal DNA | journal = Human Molecular Genetics | volume = 30 | issue = R1 | pages = R42–R48 | date = April 2021 | pmid = 33547782 | pmc = 8242491 | doi = 10.1093/hmg/ddab019 }}</ref><ref>{{Cite journal |last1=Pagani |first1=Luca |last2=Kivisild |first2=Toomas |last3=Tarekegn |first3=Ayele |last4=Ekong |first4=Rosemary |last5=Plaster |first5=Chris |last6=Gallego Romero |first6=Irene |last7=Ayub |first7=Qasim |last8=Mehdi |first8=S. Qasim |last9=Thomas |first9=Mark G. |last10=Luiselli |first10=Donata |last11=Bekele |first11=Endashaw |date=2012-07-13 |title=Ethiopian genetic diversity reveals linguistic stratification and complex influences on the Ethiopian gene pool |journal=American Journal of Human Genetics |volume=91 |issue=1 |pages=83–96 |doi=10.1016/j.ajhg.2012.05.015 |issn=1537-6605 |pmc=3397267 |pmid=22726845}}</ref>
=====Kenya=====
At Jawuoyo Rockshelter, in [[Kisumu County]], [[Kenya]], a forager of the Later [[Stone Age]] carried haplogroups [[Haplogroup E-V68#E-V22|E1b1b1a1b2/E-V22]] and [[Haplogroup L4 (mtDNA)|L4b2a2c]].<ref name="Prendergast">{{cite journal | vauthors = Prendergast ME, Lipson M, Sawchuk EA, Olalde I, Ogola CA, Rohland N, Sirak KA, Adamski N, Bernardos R, Broomandkhoshbacht N, Callan K, Culleton BJ, Eccles L, Harper TK, Lawson AM, Mah M, Oppenheimer J, Stewardson K, Zalzala F, Ambrose SH, Ayodo G, Gates HL, Gidna AO, Katongo M, Kwekason A, Mabulla AZ, Mudenda GS, Ndiema EK, Nelson C, Robertshaw P, Kennett DJ, Manthi FK, Reich D | display-authors = 6 | title = Ancient DNA reveals a multistep spread of the first herders into sub-Saharan Africa | journal = Science | volume = 365 | issue = 6448 | date = July 2019 | pmid = 31147405 | doi = 10.1126/science.aaw6275 | pmc = 6827346 | bibcode = 2019Sci...365.6275P }}</ref><ref name="Prendergast II">{{cite journal | vauthors = Prendergast ME, Lipson M, Sawchuk EA, Olalde I, Ogola CA, Rohland N, Sirak KA, Adamski N, Bernardos R, Broomandkhoshbacht N, Callan K, Culleton BJ, Eccles L, Harper TK, Lawson AM, Mah M, Oppenheimer J, Stewardson K, Zalzala F, Ambrose SH, Ayodo G, Gates HL, Gidna AO, Katongo M, Kwekason A, Mabulla AZ, Mudenda GS, Ndiema EK, Nelson C, Robertshaw P, Kennett DJ, Manthi FK, Reich D | display-authors = 6 |title=Supplementary Materials for Ancient DNA reveals a multistep spread of the first herders into sub-Saharan Africa |journal=Science | date = 5 July 2019 | volume = 365 | issue = 6448 | pages = eaaw6275 | doi = 10.1126/science.aaw6275 | pmid = 31147405 | pmc = 6827346 | bibcode = 2019Sci...365.6275P }}</ref>
At Ol Kalou, in [[Nyandarua County]], [[Kenya]], a pastoralist of the [[Pastoral Neolithic]] carried haplogroups [[Haplogroup E-Z827#E-M293|E1b1b1b2b2a1/E-M293]] and [[Haplogroup L3 (mtDNA)#Subclade distribution|L3d1d]].<ref name="Prendergast" /><ref name="Prendergast II" />
At Kokurmatakore, in [[Marsabit County]], [[Kenya]], a pastoralist of the Pastoral [[Iron Age#Sub-Saharan Africa|Iron Age]] carried haplogroups [[Haplogroup E-M35|E1b1b1/E-M35]] and [[Haplogroup L3 (mtDNA)#Subclade distribution|L3a2a]].<ref name="Prendergast" /><ref name="Prendergast II" />
At White Rock Point, in [[Homa Bay County]], [[Kenya]], there were two foragers of the Later [[Stone Age]]; one carried haplogroups [[Haplogroup BT|BT (xCT)]], likely [[Haplogroup B-M60|B]], and [[Haplogroup L2 (mtDNA)#Haplogroup L2a|L2a4]], and another probably carried [[Haplogroup L0 (mtDNA)#Distribution|haplogroup L0a2]].<ref name="Prendergast" /><ref name="Prendergast II" />
At Nyarindi Rockshelter, in [[Kenya]], there were two individuals, dated to the Later [[Stone Age]] (3500 BP); one carried [[Haplogroup L4 (mtDNA)|haplogroup L4b2a]] and another carried [[Haplogroup E-M96|haplogroup E (E-M96, E-P162)]].<ref name="Wang" /><ref name="Wang II" />
At Lukenya Hill, in [[Kenya]], there were two individuals, dated to the [[Pastoral Neolithic]] (3500 BP); one carried haplogroups [[Haplogroup E-Z827#E-V1515|E1b1b1b2b (E-M293, E-CTS10880)]] and [[Haplogroup L4 (mtDNA)|L4b2a2b]], and another carried [[Haplogroup L0 (mtDNA)#Distribution|haplogroup L0f1]].<ref name="Wang" /><ref name="Wang II" />
At Hyrax Hill, in [[Kenya]], an individual, dated to the [[Pastoral Neolithic]] (2300 BP), carried haplogroups [[Haplogroup E-Z827#E-V1515|E1b1b1b2b (E-M293, E-M293)]] and [[Haplogroup L5 (mtDNA)|L5a1b]].<ref name="Wang" /><ref name="Wang II" />
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 [[Haplogroup E-Z827#E-V1515|E1b1b1b2b (E-M293, E-M293)]] and [[Haplogroup L3 (mtDNA)#Subclade distribution|L3h1a2a1]].<ref name="Wang" /><ref name="Wang II" />
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#Sub-Saharan Africa|Iron Age]] (300 BP, 900 BP); one carried haplogroups [[Haplogroup CT|CT (CT-M168, CT-M5695)]] and [[Haplogroup L3 (mtDNA)#Subclade distribution|L3i1]], another carried [[Haplogroup L2 (mtDNA)#Haplogroup L2a1|haplogroup L2a1f]], and the last carried [[Haplogroup L2 (mtDNA)#Haplogroup L2a|haplogroup L2a5]].<ref name="Wang" /><ref name="Wang II" />
At [[Panga ya Saidi]], in [[Kenya]], an individual, estimated to date between 496 BP and 322 BP, carried haplogroups [[Haplogroup E-M215 (Y-DNA)#E-Z830 (E1b1b1b2)|E1b1b1b2]] and [[Haplogroup L4 (mtDNA)|L4b2a2]].<ref name="Skoglund">{{cite journal | vauthors = Skoglund P, Thompson JC, Prendergast ME, Mittnik A, Sirak K, Hajdinjak M, Salie T, Rohland N, Mallick S, Peltzer A, Heinze A, Olalde I, Ferry M, Harney E, Michel M, Stewardson K, Cerezo-Román JI, Chiumia C, Crowther A, Gomani-Chindebvu E, Gidna AO, Grillo KM, Helenius IT, Hellenthal G, Helm R, Horton M, López S, Mabulla AZ, Parkington J, Shipton C, Thomas MG, Tibesasa R, Welling M, Hayes VM, Kennett DJ, Ramesar R, Meyer M, Pääbo S, Patterson N, Morris AG, Boivin N, Pinhasi R, Krause J, Reich D | display-authors = 6 | title = Reconstructing Prehistoric African Population Structure | journal = Cell | volume = 171 | issue = 1 | pages = 59–71.e21 | date = September 2017 | pmid = 28938123 | pmc = 5679310 | doi = 10.1016/j.cell.2017.08.049 }}</ref>
======Laikipia County======
At Kisima Farm/Porcupine Cave, in [[Laikipia County]], [[Kenya]], there were two pastoralists of the [[Pastoral Neolithic]]; one carried haplogroups [[Haplogroup E-Z827#E-M293|E1b1b1b2b2a1/E-M293]] and [[Haplogroup M (mtDNA)#Haplogroup M1|M1a1]], and another carried haplogroup [[Haplogroup M (mtDNA)#Haplogroup M1|M1a1f]].<ref name="Prendergast" /><ref name="Prendergast II" />
At Kisima Farm/C4, in [[Laikipia County]], [[Kenya]], a pastoralist of the Pastoral [[Iron Age#Sub-Saharan Africa|Iron Age]], carried haplogroups [[Haplogroup E-M75|E2 (xE2b)/E-M75]] and [[Haplogroup L3 (mtDNA)#Subclade distribution|L3h1a1]].<ref name="Prendergast" /><ref name="Prendergast II" />
At Laikipia District Burial, in [[Laikipia County]], [[Kenya]], a pastoralist of the Pastoral [[Iron Age#Sub-Saharan Africa|Iron Age]] carried haplogroup [[Haplogroup L0 (mtDNA)#Distribution|L0a1c1]].<ref name="Prendergast" /><ref name="Prendergast II" />
======Nakuru County======
At Prettejohn's Gully, in [[Nakuru County, Kenya]], there were two pastoralists of the early pastoral period; one carried haplogroups [[Haplogroup E-M75|E2 (xE2b)/E-M75]] and [[Haplogroup K (mtDNA)|K1a]], and another carried haplogroup [[Haplogroup L3 (mtDNA)#Subclade distribution|L3f1b]].<ref name="Prendergast" /><ref name="Prendergast II" />
At Cole's Burial, in [[Nakuru County, Kenya]], a pastoralist of the [[Pastoral Neolithic]] carried haplogroups [[Haplogroup E-V68#Subclades of M78|E1b1b1a1a1b1/E-CTS3282]] and [[Haplogroup L3 (mtDNA)#Subclade distribution|L3i2]].<ref name="Prendergast" /><ref name="Prendergast II" />
At Rigo Cave, in [[Nakuru County, Kenya]], there were three pastoralists of the [[Pastoral Neolithic]]/[[Elmenteitan]], one carried haplogroups [[Haplogroup E-Z827#E-M293|E1b1b1b2b2a1/E-M293]] and [[Haplogroup L3 (mtDNA)#Subclade distribution|L3f]], another carried haplogroups [[Haplogroup E-Z827#E-V1515|E1b1b1b2b2/E-V1486]], likely [[Haplogroup E-M215 (Y-DNA)#E-M293|E-M293]], and probably [[Haplogroup M (mtDNA)#Haplogroup M1|M1a1b]], and the last carried haplogroups [[Haplogroup E-M215 (Y-DNA)#E-M293|E1b1b1b2b2a1/E-M293]] and [[Haplogroup L4 (mtDNA)|L4b2a2c]].<ref name="Prendergast" /><ref name="Prendergast II" />
At Naishi Rockshelter, in [[Nakuru County, Kenya]], there two pastoralists of the [[Pastoral Neolithic]]; one carried haplogroups [[Haplogroup E-Z827#E-V1515|E1b1b1b2b/E-V1515]], likely [[Haplogroup E-M215 (Y-DNA)#E-M293|E-M293]], and [[Haplogroup L3 (mtDNA)#Subclade distribution|L3x1a]], and another carried haplogroups [[Haplogroup A (Y-DNA)#Structure & subclade distribution|A1b (xA1b1b2a)/A-P108]] and [[Haplogroup L0 (mtDNA)#Distribution|L0a2d]].<ref name="Prendergast" /><ref name="Prendergast II" />
At Keringet Cave, in [[Nakuru County, Kenya]], a pastoralist of the [[Pastoral Neolithic]] carried [[Haplogroup A (Y-DNA)#Structure & subclade distribution|haplogroups A1b1b2/A-L427]] and [[Haplogroup L4 (mtDNA)|L4b2a1]], and another pastoralist of the [[Pastoral Neolithic]]/[[Elmenteitan]] carried [[Haplogroup K (mtDNA)|haplogroup K1a]].<ref name="Prendergast" /><ref name="Prendergast II" />
At Naivasha Burial Site, in [[Nakuru County, Kenya]], there were five pastoralists of the [[Pastoral Neolithic]]; one carried [[Haplogroup L4 (mtDNA)|haplogroup L4b2a2b]], another carried haplogroups xBT, likely [[Haplogroup A (Y-DNA)|A]], and [[Haplogroup M (mtDNA)#Haplogroup M1|M1a1b]], another carried haplogroups [[Haplogroup E-Z827#E-M293|E1b1b1b2b2a1/E-M293]] and [[Haplogroup L3 (mtDNA)#Subclade distribution|L3h1a1]], another carried haplogroups [[Haplogroup A (Y-DNA)#A1b1b2b (A-M13)|A1b1b2b/A-M13]] and [[Haplogroup L4 (mtDNA)|L4a1]], and the last carried haplogroups [[Haplogroup E-Z827#E-M293|E1b1b1b2b2a1/E-M293]] and [[Haplogroup L3 (mtDNA)#Subclade distribution|L3x1a]].<ref name="Prendergast" /><ref name="Prendergast II" />
At Njoro River Cave II, in [[Nakuru County, Kenya]], a pastoralist of the [[Pastoral Neolithic]] carried haplogroup [[Haplogroup L3 (mtDNA)#Subclade distribution|L3h1a2a1]].<ref name="Prendergast" /><ref name="Prendergast II" />
At Egerton Cave, in [[Nakuru County, Kenya]], a pastoralist of the [[Pastoral Neolithic]]/[[Elmenteitan]] carried haplogroup [[Haplogroup L0 (mtDNA)#Distribution|L0a1d]].<ref name="Prendergast" /><ref name="Prendergast II" />
At Ilkek Mounds, in [[Nakuru County, Kenya]], a pastoralist of the Pastoral [[Iron Age#Sub-Saharan Africa|Iron Age]] carried haplogroups [[Haplogroup E-M75|E2 (xE2b)/E-M75]] and [[Haplogroup L0 (mtDNA)#Distribution|L0f2a]].<ref name="Prendergast" /><ref name="Prendergast II" />
At Deloraine Farm, in [[Nakuru County, Kenya]], an iron metallurgist of the [[Iron Age#Sub-Saharan Africa|Iron Age]] carried haplogroups [[Haplogroup E-M2#E1b1a1a1a|E1b1a1a1a1a/E-M58]] and [[Haplogroup L5 (mtDNA)|L5b1]].<ref name="Prendergast" /><ref name="Prendergast II" />
======Narok County======
At Kasiole 2, in [[Narok County]], [[Kenya]], a pastoralist of the Pastoral [[Iron Age#Sub-Saharan Africa|Iron Age]] carried haplogroups [[Haplogroup E-Z827#E-V1515|E1b1b1b2b/E-V1515]], likely [[Haplogroup E-Z827#E-M293|E-M293]], and [[Haplogroup L3 (mtDNA)#Subclade distribution|L3h1a2a1]].<ref name="Prendergast" /><ref name="Prendergast II" />
At Emurua Ole Polos, in [[Narok County]], [[Kenya]], a pastoralist of the Pastoral [[Iron Age#Sub-Saharan Africa|Iron Age]] carried haplogroups [[Haplogroup E-Z827#E-M293|E1b1b1b2b2a1/E-M293]] and [[Haplogroup L3 (mtDNA)#Subclade distribution|L3h1a2a1]].<ref name="Prendergast" /><ref name="Prendergast II" />
=====Tanzania=====
At [[Luxmanda]], [[Tanzania]], an individual, estimated to date between 3141 BP and 2890 BP, carried [[Haplogroup L2 (mtDNA)#Haplogroup L2a1|haplogroup L2a1]].<ref name="Skoglund" />
At [[Kuumbi Cave, Zanzibar|Kuumbi Cave]], in [[Zanzibar]], [[Tanzania]], an individual, estimated to date between 1370 BP and 1303 BP, carried haplogroup [[Haplogroup L4 (mtDNA)|L4b2a2c]].<ref name="Skoglund" />
======Karatu District======
At Gishimangeda Cave, in [[Karatu District]], [[Tanzania]], there were eleven pastoralists of the [[Pastoral Neolithic]]; one carried haplogroups [[Haplogroup E-V68#Family tree|E1b1b1a1b2/E-V22]] and [[Haplogroup HV (mtDNA)#Tree|HV1b1]], another carried [[Haplogroup L0 (mtDNA)#Distribution|haplogroup L0a]], another carried haplogroup [[Haplogroup L3 (mtDNA)#Subclade distribution|L3x1]], another carried haplogroup [[Haplogroup L4 (mtDNA)|L4b2a2b]], another carried haplogroups [[Haplogroup E-Z827#E-M293|E1b1b1b2b2a1/E-M293]] and [[Haplogroup L3 (mtDNA)#Subclade distribution|L3i2]], another carried haplogroup [[Haplogroup L3 (mtDNA)#Subclade distribution|L3h1a2a1]], another carried haplogroups [[Haplogroup E-Z827#E-V1515|E1b1b1b2b2/E-V1486]], likely [[Haplogroup E-Z827#E-M293|E-M293]] and [[Haplogroup L0 (mtDNA)#Distribution|L0f2a1]], and another carried haplogroups [[Haplogroup E-Z827#E-V1515|E1b1b1b2b2/E-V1486]], likely [[Haplogroup E-Z827#E-M293|E-M293]], and [[Haplogroup T (mtDNA)|T2+150]]; while most of the haplogroups among three pastoralists went undetermined, one was determined to carry [[haplogroup BT]], likely [[Haplogroup B-M60|B]].<ref name="Prendergast" /><ref name="Prendergast II" />
======Pemba Island======
At Makangale Cave, on [[Pemba Island]], [[Tanzania]], an individual, estimated to date between 1421 BP and 1307 BP, carried [[Haplogroup L0 (mtDNA)#Distribution|haplogroup L0a]].<ref name="Skoglund" />
At Makangale Cave, on [[Pemba Island]], [[Tanzania]], an individual, estimated to date between 639 BP and 544 BP, carried [[Haplogroup L2 (mtDNA)#Haplogroup L2a1a|haplogroup L2a1a2]].<ref name="Skoglund" />
=====Uganda=====
At [[Munsa]], in [[Uganda]], an individual, dated to the Later [[Iron Age#Sub-Saharan Africa|Iron Age]] (500 BP), carried haplogroup [[Haplogroup L3 (mtDNA)#Subclade distribution|L3b1a1]].<ref name="Wang" /><ref name="Wang II" />
====Y-Chromosomal DNA====
{{Further|Y-DNA haplogroups in populations of Sub-Saharan Africa}}
As of 19,000 years ago, Africans, bearing [[Haplogroup E-V38|haplogroup E1b1a-V38]], likely traversed across the [[Sahara]], from [[East Africa|east]] to [[West Africa|west]].<ref name="Shriner"/>
Before the [[Trans-Saharan slave trade#Trans-Saharan slave trade in the Middle Ages|slave]] [[Indian Ocean slave trade#Muslim Period|trade]] period, East Africans, who carried [[Haplogroup E-M2|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]]).<ref name="Fernandes">{{cite web | vauthors = Neves da Nova Fernandes VC |title=High-resolution characterization of genetic markers in the Arabian Peninsula and Near East |url=https://etheses.whiterose.ac.uk/6301/1/Vfernandes_PhD_thesis.pdf |website=White Rose eTheses Online |publisher=University of Leeds}}</ref>
====Mitochondrial DNA====
{{Further|Macro-haplogroup L (mtDNA)|Haplogroup M (mtDNA)|Haplogroup N (mtDNA)}}
====Autosomal DNA====
Across all areas of [[Madagascar]], the average ancestry for the [[Malagasy people]] was found to be 4% [[West Eurasian]], 37% [[Austronesian peoples|Austronesian]], and 59% [[Bantu peoples|Bantu]].<ref name="Heiske">{{cite journal | vauthors = Heiske M, Alva O, Pereda-Loth V, Van Schalkwyk M, Radimilahy C, Letellier T, Rakotarisoa JA, Pierron D | display-authors = 6 | title = Genetic evidence and historical theories of the Asian and African origins of the present Malagasy population | journal = Human Molecular Genetics | volume = 30 | issue = R1 | pages = R72–R78 | date = April 2021 | pmid = 33481023 | doi = 10.1093/hmg/ddab018 | doi-access = free }}</ref>
====Medical DNA====
===Southern Africa===
{{See also|San people#Genetics|Khoekhoe}}
====Archaic Human DNA====
{{Further|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.<ref name="Bergström"/>
====Ancient DNA====
Three [[Later Stone Age]] [[hunter-gatherers]] carried ancient DNA similar to [[Khoisan]]-speaking hunter-gatherers.<ref name="Choudhury">{{cite journal | vauthors = Choudhury A, Sengupta D, Ramsay M, Schlebusch C | title = Bantu-speaker migration and admixture in southern Africa | journal = Human Molecular Genetics | volume = 30 | issue = R1 | pages = R56–R63 | date = April 2021 | pmid = 33367711 | pmc = 8117461 | doi = 10.1093/hmg/ddaa274 }}</ref> Prior to the [[Bantu migration]] into the region, as evidenced by ancient DNA from [[Botswana]], [[East Africa]]n [[herders]] migrated into Southern Africa.<ref name="Choudhury" /> Out of four [[Iron Age]] Bantu [[agriculturalists]] of [[West African]] origin, two earlier agriculturalists carried ancient DNA similar to [[Tsonga people|Tsonga]] and [[Venda people|Venda]] peoples and the two later agriculturalists carried ancient DNA similar to [[Nguni people]]; this indicates that there were various movements of peoples in the overall Bantu migration, which resulted in increased interaction and admixing between [[Bantu languages|Bantu-speaking]] peoples and Khoisan-speaking peoples.<ref name="Choudhury" />
=====Botswana=====
At Nqoma, in [[Botswana]], an individual, dated to the Early [[Iron Age#Sub-Saharan Africa|Iron Age]] (900 BP), carried [[Haplogroup L2 (mtDNA)#Haplogroup L2a1|haplogroup L2a1f]].<ref name="Wang" /><ref name="Wang II" />
At Taukome, in [[Botswana]], an individual, dated to the Early [[Iron Age#Sub-Saharan Africa|Iron Age]] (1100 BP), carried haplogroups [[Haplogroup E-M2#E1b1a1|E1b1a1 (E-M2, E-Z1123)]] and [[Haplogroup L0 (mtDNA)#Distribution|L0d3b1]].<ref name="Wang" /><ref name="Wang II" />
At Xaro, in [[Botswana]], there were two individuals, dated to the Early [[Iron Age#Sub-Saharan Africa|Iron Age]] (1400 BP); one carried haplogroups [[Haplogroup E-M2#E1b1a1a1c|E1b1a1a1c1a]] and [[Haplogroup L3 (mtDNA)#Subclade distribution|L3e1a2]], and another carried haplogroups [[Haplogroup E-Z827#E-V1515|E1b1b1b2b (E-M293, E-CTS10880)]] and [[Haplogroup L0 (mtDNA)#Distribution|L0k1a2]].<ref name="Wang" /><ref name="Wang II" />
=====Malawi=====
======Fingira======
At Fingira, in [[Malawi]], an individual, estimated to date between 6175 BP and 5913 BP, carried haplogroups [[Haplogroup BT|BT]] and [[Haplogroup L0 (mtDNA)#Distribution|L0d1b2b]].<ref name="Skoglund" />
At Fingira, in [[Malawi]], an individual, estimated to date between 6177 BP and 5923 BP, carried haplogroups [[Haplogroup BT|BT]] and [[Haplogroup L0 (mtDNA)#Distribution|L0d1c]].<ref name="Skoglund" />
At Fingira, in [[Malawi]], an individual, estimated to date between 2676 BP and 2330 BP, carried [[Haplogroup L0 (mtDNA)#Distribution|haplogroup L0f]].<ref name="Skoglund" />
======Chencherere======
At Chencherere, in [[Malawi]], an individual, estimated to date between 5400 BP and 4800 BP, carried [[Haplogroup L0 (mtDNA)#Distribution|haplogroup L0k2]].<ref name="Skoglund" />
At Chencherere, in [[Malawi]], an individual, estimated to date between 5293 BP and 4979 BP, carried [[Haplogroup L0 (mtDNA)#Distribution|haplogroup L0k1]].<ref name="Skoglund" />
======Hora======
At Hora, in [[Malawi]], an individual, estimated to date between 10,000 BP and 5000 BP, carried haplogroups [[Haplogroup BT|BT]] and [[Haplogroup L0 (mtDNA)#Distribution|L0k2]].<ref name="Skoglund" />
At Hora, in [[Malawi]], an individual, estimated to date between 8173 BP and 7957 BP, carried [[Haplogroup L0 (mtDNA)#Distribution|haplogroup L0a2]].<ref name="Skoglund" />
=====South Africa=====
At [[Doonside, KwaZulu-Natal|Doonside]], in [[South Africa]], an individual, estimated to date between 2296 BP and 1910 BP, carried [[Haplogroup L0 (mtDNA)#Distribution|haplogroup L0d2]].<ref name="Schlebusch" /><ref name="Schlebusch II" />
At [[Champagne Castle]], in [[South Africa]], an individual, estimated to date between 448 BP and 282 BP, carried [[Haplogroup L0 (mtDNA)#Distribution|haplogroup L0d2a1a]].<ref name="Schlebusch" /><ref name="Schlebusch II" />
At [[Elands Bay Cave|Eland Cave]], in [[South Africa]], an individual, estimated to date between 533 BP and 453 BP, carried [[Haplogroup L3 (mtDNA)#Subclade distribution|haplogroup L3e3b1]].<ref name="Schlebusch" /><ref name="Schlebusch II" />
At Mfongosi, in [[South Africa]], an individual, estimated to date between 448 BP and 308 BP, carried [[Haplogroup L3 (mtDNA)#Subclade distribution|haplogroup L3e1b2]].<ref name="Schlebusch" /><ref name="Schlebusch II" />
At [[Newcastle, KwaZulu-Natal|Newcastle]], in [[South Africa]], an individual, estimated to date between 508 BP and 327 BP, carried [[Haplogroup L3 (mtDNA)#Subclade distribution|haplogroup L3e2b1a2]].<ref name="Schlebusch" /><ref name="Schlebusch II" />
At [[St Helena Bay|St. Helena]], in [[South Africa]], an individual, estimated to date between 2241 BP and 1965 BP, carried haplogroups [[Haplogroup A (Y-DNA)#A1b1b2a (A-M51)|A1b1b2a]] and [[Haplogroup L0 (mtDNA)#Distribution|L0d2c1]].<ref name="Skoglund" />
At Faraoskop Rock Shelter, in [[South Africa]], an individual, estimated to date between 2017 BP and 1748 BP, carried haplogroups [[Haplogroup A (Y-DNA)#A1b1b2a (A-M51)|A1b1b2a]] and [[Haplogroup L0 (mtDNA)#Distribution|L0d1b2b1b]].<ref name="Skoglund" />
At Kasteelberg, in [[South Africa]], an individual, estimated to date between 1282 BP and 1069 BP, carried haplogroup [[Haplogroup L0 (mtDNA)#Distribution|L0d1a1a]].<ref name="Skoglund" />
At Vaalkrans Shelter, in [[South Africa]], an individual, estimated to date to 200 BP, is predominantly related to [[Khoisan]] speakers, partly related (15% - 32%) to [[East Africa#Demographics|East Africans]], and carried haplogroups [[Haplogroup L0 (mtDNA)#Distribution|L0d3b1]].<ref name="Coutinho">{{cite journal | vauthors = Coutinho A, Malmström H, Edlund H, Henshilwood CS, van Niekerk KL, Lombard M, Schlebusch CM, Jakobsson M | display-authors = 6 | title = Later Stone Age human hair from Vaalkrans Shelter, Cape Floristic Region of South Africa, reveals genetic affinity to Khoe groups | journal = American Journal of Physical Anthropology | volume = 174 | issue = 4 | pages = 701–713 | date = April 2021 | pmid = 33539553 | doi = 10.1002/ajpa.24236 | publisher = Am J Phys Anthropol | s2cid = 213563734 }}</ref>
======Ballito Bay======
At [[Ballito]] Bay, [[South Africa]], an individual, estimated to date between 1986 BP and 1831 BP, carried haplogroups [[Haplogroup A (Y-DNA)#Structure & subclade distribution|A1b1b2]] and [[Haplogroup L0 (mtDNA)#Distribution|L0d2c1]].<ref name="Schlebusch">{{cite journal | vauthors = Schlebusch CM, Malmström H, Günther T, Sjödin P, Coutinho A, Edlund H, Munters AR, Vicente M, Steyn M, Soodyall H, Lombard M, Jakobsson M | display-authors = 6 | title = Southern African ancient genomes estimate modern human divergence to 350,000 to 260,000 years ago | journal = Science | volume = 358 | issue = 6363 | pages = 652–655 | date = November 2017 | pmid = 28971970 | doi = 10.1126/science.aao6266 | s2cid = 206663925 | doi-access = free | bibcode = 2017Sci...358..652S }}</ref><ref name="Schlebusch II">{{cite journal | vauthors = Schlebusch CM, Malmström H, Günther T, Sjödin P, Coutinho A, Edlund H, Munters AR, Vicente M, Steyn M, Soodyall H, Lombard M, Jakobsson M | display-authors = 6 |title=Supplementary Materials for Southern African ancient genomes estimate modern human divergence to 350,000to 260,000years ago |url=https://www.science.org/doi/10.1126/science.aao6266 |journal=Science | date = 3 November 2017 | volume = 358 | issue = 6363 | pages = 652–655 | doi = 10.1126/science.aao6266 | pmid = 28971970 | bibcode = 2017Sci...358..652S | s2cid = 206663925 }}</ref>
At [[Ballito]] Bay, [[South Africa]], an individual, estimated to date between 2149 BP and 1932 BP, carried haplogroups [[Haplogroup A (Y-DNA)#Structure & subclade distribution|A1b1b2]] and [[Haplogroup L0 (mtDNA)#Distribution|L0d2a1]].<ref name="Schlebusch" /><ref name="Schlebusch II" />
====Y-Chromosomal DNA====
{{Further|Y-DNA haplogroups in populations of Sub-Saharan Africa}}
[[File:Distribution of Y-Chromosome Haplogroup A in Africa.png|thumb|Distribution of Y-Chromosome Haplogroup A in Africa.]]
[[File:B haplogroup of Y-DNA.png|thumb|Distribution of haplogroup B (M60) of the human Y chromosome in native populations.]]
Various [[Y chromosome]] studies show that the San carry some of the most divergent (oldest) [[Human Y-chromosome DNA haplogroup|human Y-chromosome haplogroup]]s. These haplogroups are specific sub-groups of haplogroups [[Haplogroup A (Y-DNA)|A]] and [[Haplogroup B (Y-DNA)|B]], the two earliest branches on the human Y-chromosome [[phylogenetic tree|tree]].<ref name=j1>{{cite journal | vauthors = Knight A, Underhill PA, Mortensen HM, Zhivotovsky LA, Lin AA, Henn BM, Louis D, Ruhlen M, Mountain JL | display-authors = 6 | title = African Y chromosome and mtDNA divergence provides insight into the history of click languages | journal = Current Biology | volume = 13 | issue = 6 | pages = 464–473 | date = March 2003 | pmid = 12646128 | doi = 10.1016/S0960-9822(03)00130-1 | s2cid = 52862939 | doi-access = free }}</ref><ref>{{cite journal | vauthors = Hammer MF, Karafet TM, Redd AJ, Jarjanazi H, Santachiara-Benerecetti S, Soodyall H, Zegura SL | title = Hierarchical patterns of global human Y-chromosome diversity | journal = Molecular Biology and Evolution | volume = 18 | issue = 7 | pages = 1189–1203 | date = July 2001 | pmid = 11420360 | doi = 10.1093/oxfordjournals.molbev.a003906 | doi-access = free }}</ref><ref>{{cite journal | vauthors = Naidoo T, Schlebusch CM, Makkan H, Patel P, Mahabeer R, Erasmus JC, Soodyall H | title = Development of a single base extension method to resolve Y chromosome haplogroups in sub-Saharan African populations | journal = Investigative Genetics | volume = 1 | issue = 1 | pages = 6 | date = September 2010 | pmid = 21092339 | pmc = 2988483 | doi = 10.1186/2041-2223-1-6 }}</ref>
====Mitochondrial DNA====
[[Mitochondrial DNA]] studies also provide evidence that the San carry high frequencies of the earliest [[Human mitochondrial DNA haplogroup|haplogroup]] branches in the human mitochondrial DNA tree. This DNA is inherited only from one's mother. The most divergent (oldest) mitochondrial haplogroup, [[Haplogroup L0 (mtDNA)|L0]]d, has been identified at its highest frequencies in the southern African San groups.<ref name=j1/><ref>{{cite journal | vauthors = Chen YS, Olckers A, Schurr TG, Kogelnik AM, Huoponen K, Wallace DC | title = mtDNA variation in the South African Kung and Khwe-and their genetic relationships to other African populations | journal = American Journal of Human Genetics | volume = 66 | issue = 4 | pages = 1362–1383 | date = April 2000 | pmid = 10739760 | pmc = 1288201 | doi = 10.1086/302848 }}</ref><ref>{{cite journal | vauthors = Tishkoff SA, Gonder MK, Henn BM, Mortensen H, Knight A, Gignoux C, Fernandopulle N, Lema G, Nyambo TB, Ramakrishnan U, Reed FA, Mountain JL | display-authors = 6 | title = History of click-speaking populations of Africa inferred from mtDNA and Y chromosome genetic variation | journal = Molecular Biology and Evolution | volume = 24 | issue = 10 | pages = 2180–2195 | date = October 2007 | pmid = 17656633 | doi = 10.1093/molbev/msm155 | doi-access = free }}</ref><ref>{{cite journal | vauthors = Schlebusch CM, Naidoo T, Soodyall H | title = SNaPshot minisequencing to resolve mitochondrial macro-haplogroups found in Africa | journal = Electrophoresis | volume = 30 | issue = 21 | pages = 3657–3664 | date = November 2009 | pmid = 19810027 | doi = 10.1002/elps.200900197 | s2cid = 19515426 }}</ref>
====Autosomal DNA====
In a study published in March 2011, Brenna Henn and colleagues found that the ǂKhomani San, as well as the [[Sandawe people|Sandawe]] and [[Hadza people]]s of [[Tanzania]], were the most genetically diverse of any living humans studied. This high degree of genetic diversity hints at the origin of [[anatomically modern humans]].<ref>{{cite journal | vauthors = Henn BM, Gignoux CR, Jobin M, Granka JM, Macpherson JM, Kidd JM, Rodríguez-Botigué L, Ramachandran S, Hon L, Brisbin A, Lin AA, Underhill PA, Comas D, Kidd KK, Norman PJ, Parham P, Bustamante CD, Mountain JL, Feldman MW | display-authors = 6 | title = Hunter-gatherer genomic diversity suggests a southern African origin for modern humans | journal = Proceedings of the National Academy of Sciences of the United States of America | volume = 108 | issue = 13 | pages = 5154–5162 | date = March 2011 | pmid = 21383195 | pmc = 3069156 | doi = 10.1073/pnas.1017511108 | publisher = [[National Academy of Sciences]] | doi-access = free }}</ref><ref>{{cite journal |last=Kaplan |first=Matt |year=2011 |title=Gene Study Challenges Human Origins in Eastern Africa |journal=[[Scientific American]]|publisher=[[Nature Publishing Group]] |url=http://www.scientificamerican.com/article/gene-study-challenges-human-origin-africa/ |access-date=22 June 2012}}</ref>
====Medical DNA====
Among the ancient DNA from three [[hunter-gatherers]] sharing genetic similarity with [[San people]] and four Iron Age [[agriculturalists]], their [[SNPs]] indicated that they bore variants for resistance against [[sleeping sickness]] and [[Plasmodium vivax]].<ref name="Pfeiffer">{{cite journal | vauthors = Pfeiffer S | title = Disease as a Factor in the African Archaeological Record | journal = The African Archaeological Review | volume = 37 | issue = 3 | pages = 487–490 | year = 2020 | pmid = 32863518 | pmc = 7445818 | doi = 10.1007/s10437-020-09405-7 }}</ref> In particular, two out of the four Iron Age agriculturalists bore variants for resistance against sleeping sickness and three out of the four Iron Age agriculturalists bore [[Human genetic resistance to malaria#Duffy antigen receptor negativity|Duffy negative variants]] for resistance against [[malaria]].<ref name="Pfeiffer" /> In contrast to the Iron Age agriculturalists, from among the San-related hunter-gatherers, a six-year-old boy may have died from [[schistosomiasis]].<ref name="Pfeiffer" /> In [[Botswana]], a man, who dates to 1400 BP, may have also carried the Duffy negative variant for resistance against malaria.<ref name="Pfeiffer" />
==Recent African origin of modern humans==
{{Main|Recent African origin of modern humans}}
{{Further|Early human migrations|Basal Eurasian|Human genetic variation}}
Between 500,000 BP and 300,000 BP, [[anatomically modern humans]] may have emerged in Africa.<ref name="Pereira">{{cite journal | vauthors = Pereira L, Mutesa L, Tindana P, Ramsay M | title = African genetic diversity and adaptation inform a precision medicine agenda | journal = Nature Reviews. Genetics | volume = 22 | issue = 5 | pages = 284–306 | date = May 2021 | pmid = 33432191 | doi = 10.1038/s41576-020-00306-8 | publisher = Nature Reviews | s2cid = 231587564 }}</ref> As Africans (e.g., [[Y-Chromosomal Adam]], [[Mitochondrial Eve]]) have migrated from their places of origin in Africa to other locations in Africa, and as the time of divergence for [[East Africa]]n, [[Central Africa]]n, and [[West African]] lineages are similar to the time of divergence for the [[Southern African]] lineage, there is insufficient evidence to identify a specific region for the origin of humans in [[Africa]].<ref name="Bergström" /> In 100,000 BP, anatomically modern humans migrated from [[Africa]] into [[Eurasia]].<ref name="Benton">{{cite journal | vauthors = Benton ML, Abraham A, LaBella AL, Abbot P, Rokas A, Capra JA | title = The influence of evolutionary history on human health and disease | journal = Nature Reviews. Genetics | volume = 22 | issue = 5 | pages = 269–283 | date = May 2021 | pmid = 33408383 | pmc = 7787134 | doi = 10.1038/s41576-020-00305-9 }}</ref> Subsequently, tens of thousands of years after, the ancestors of all present-day Eurasians migrated from Africa into Eurasia and eventually became admixed with [[Denisovans]] and [[Neanderthals]].<ref name="Benton" />
== See also ==
*[[Genetic history of the African diaspora]]
== Notes ==
{{NoteFoot}}
== References ==
{{reflist}}
{{Human genetics}}
[[Category:Genetic genealogy]]
[[Category:History of Africa]]
[[Category:Modern human genetic history]]' |
New page wikitext, after the edit (new_wikitext) | '[[File:ADMIXTURE analysis of Horn of Africa populations in a broad context.png|thumb|Population structure of African populations in a broad context. ADMIXTURE analysis of 2,194 individuals from 81 populations for 16,420 SNPs reveals both well-established and novel ancestry components in African populations.]]
The '''genetic history of Africa''' is composed of the overall [[genetic history]] of [[Africa|African populations]], including the regional genetic histories of [[North Africa]], [[West Africa]], [[East Africa]], [[Central Africa]], and [[Southern Africa]], as well as the [[recent African origin of modern humans|recent origin of modern humans in Africa]]. The [[Sahara]] served as a trans-regional passageway and place of dwelling for people in Africa during various [[Interstadial|humid phases]]<ref name="Osborn">{{cite journal |vauthors=Osborne AH, Vance D, Rohling EJ, Barton N, Rogerson M, Fello N |title=A humid corridor across the Sahara for the migration of early modern humans out of Africa 120,000 years ago |journal=Proceedings of the National Academy of Sciences of the United States of America |volume=105 |issue=43 |pages=16444–16447 |date=October 2008 |pmid=18936490 |pmc=2575439 |doi=10.1073/pnas.0804472105 |s2cid=10418009 |doi-access=free |bibcode=2008PNAS..10516444O}}</ref><ref name="Drake">{{cite book |vauthors=Drake N, Breeze P |title=Africa from MIS 6-2 |chapter=Climate Change and Modern Human Occupation of the Sahara from MIS 6-2 |chapter-url=https://link.springer.com/chapter/10.1007/978-94-017-7520-5_6 |website=SpringerLink |series=Vertebrate Paleobiology and Paleoanthropology |year=2016 |pages=103–122 |publisher=Africa from MIS 6-2 |doi=10.1007/978-94-017-7520-5_6 |isbn=978-94-017-7519-9}}</ref><ref name="El-Shenawy">{{cite journal |vauthors=El-Shenawy MI, Kim ST, Schwarcz HP, Asmerom Y, Polyak VJ |title=Speleothem evidence for the greening of the Sahara and its implications for the early human dispersal out of sub-Saharan Africa |url=https://www.sciencedirect.com/science/article/abs/pii/S0277379117307436 |journal=Quaternary Science Reviews |year=2018 |volume=188 |pages=67–76 |doi=10.1016/j.quascirev.2018.03.016 |bibcode=2018QSRv..188...67E}}</ref> and periods throughout the [[history of Africa]].<ref name="Scheele">{{cite journal |vauthors=Scheele J |title=Crossroads Regions: The Sahara |url=https://www.academia.edu/37787505 |website=Academia |publisher=Oxford Handbooks Online}}</ref><ref name="Wippel">{{cite journal |vauthors=Wippel S |title=The Sahara as a Bridge, Not a Barrier: An Essay and Book Review on Recent Transregional Perspectives |journal=Neue Politische Literatur |year=2020 |volume=65 |issue=3 |pages=449–472 |doi=10.1007/s42520-020-00318-y |doi-access=free}}</ref>
==Overview==
[[File:PCA of sub-Saharan Africa populations.png|thumb|(A) the origin of the 46 African ethnic groups used in the analysis; ethnic groups from similar regions are given the same colour, but different shapes. (B) PCA shows that the first major axis of variation in Africa (PC1, y-axis) splits southern groups from the rest of Africa, each symbol represents an individual; PC2 (x-axis) reflects ethno-linguistic differences, with Niger-Congo and Nilo-Saharan speakers split from Afroasiatic speakers. (C) The third principle component (PC3, x-axis) represents geographical separation of Niger-Congo speakers, forming a cline from west to east Africans.]]
The people of [[Africa]] are characterized by regional genetic substructure and heterogeneity, depending on the respective ethno-linguistic identity, and, in part, explainable by the "multiregional evolution" of modern human lineages in various multiple regions of the African continent, as well as later admixture events, including back-migrations from Eurasia, of both highly differentiated West- and East-Eurasian components.<ref name="The deep population history in Afri">{{cite journal |vauthors=Hollfelder N, Breton G, Sjödin P, Jakobsson M |title=The deep population history in Africa |journal=Human Molecular Genetics |volume=30 |issue=R1 |pages=R2–R10 |date=April 2021 |pmid=33438014 |pmc=8117439 |doi=10.1093/hmg/ddab005}}</ref>
Africans' genetic ancestry is largely partitioned by [[geography]] and [[language family]], with populations belonging to the same ethno-linguistic groupings showing high genetic homogeneity and coherence. Gene flow, consistent with both short- and long-range migration events followed by extensive admixture and [[bottleneck effect|bottleneck event]]s, have influenced the regional genetic makeup and demographic structure of Africans. The historical [[Bantu expansion]] had lasting impacts on the modern demographic make up of Africa, resulting in a greater genetic and linguistic homogenization.<ref name="Fan 82">{{cite journal |vauthors=Fan S, Kelly DE, Beltrame MH, Hansen ME, Mallick S, Ranciaro A, Hirbo J, Thompson S, Beggs W, Nyambo T, Omar SA, Meskel DW, Belay G, Froment A, Patterson N, Reich D, Tishkoff SA |display-authors=6 |title=African evolutionary history inferred from whole genome sequence data of 44 indigenous African populations |journal=Genome Biology |volume=20 |issue=1 |pages=82 |date=April 2019 |pmid=31023338 |pmc=6485071 |doi=10.1186/s13059-019-1679-2}}</ref> Genetic, archeologic, and linguistic studies added extra insight into this movement: "Our results reveal a genetic continuum of Niger–Congo speaker populations across the continent and extend our current understanding of the routes, timing and extent of the Bantu migration."<ref>{{Cite web |title=A great African gene migration |url=https://cosmosmagazine.com/science/a-great-migration-of-genes-in-africa/ |access-date=2022-01-11 |website=cosmosmagazine.com |date=29 October 2020 |language=en-AU}}</ref>
Overall, different African populations display genetic diversity and substructure, but can be clustered in distinct but partially overlapping groupings:<ref name=":5">{{Cite web |last=Ananyo Choudhury, Shaun Aron, Dhriti Sengupta, Scott Hazelhurst, Michèle Ramsay |date=1 August 2018 |title=African genetic diversity provides novel insights into evolutionary history and local adaptations |url=https://academic.oup.com/hmg/article/27/R2/R209/4993963 |access-date=2022-10-21 |website=academic.oup.com}}</ref>
* [[Khoisan]] hunter-gatherer lineages from [[Southern Africa]] represent the deepest lineages, forming a divergent and distinct cluster, shifted away from contemporary "Sub-Saharan Africans", and are as diverged from them as the various Eurasian lineages are. The diverging date of these "Southern hunter-gatherers" from all other human populations is estimated to over 100,000 years ago respectively, with the Khoisan later diverging into two subgroups, northern and southern Khoisan,~30,000 years ago.{{NoteTag|Although the estimated date of this event varies across studies, there is general consensus that the split occurred over 100 thousand years ago (kya) (8,23).}}
* Rain forest foragers such as the [[Baka people (Cameroon and Gabon)|Baka]] and the [[Mbuti people|Mbuti]] diverged from other Sub-Saharan African groups over 60,000 years ago. Eastern groups such as the Mbuti split from Western groups such as the Baka ~20,000 years ago.
* [[Proto-Afroasiatic language|Proto-Afroasiatic]] speakers are suggested to have diverged from other African groups ~50,000 years ago.<ref>{{Cite journal |last=Baker |first=Jennifer L. |last2=Rotimi |first2=Charles N. |last3=Shriner |first3=Daniel |date=2017-05-08 |title=Human ancestry correlates with language and reveals that race is not an objective genomic classifier |url=https://www.nature.com/articles/s41598-017-01837-7 |journal=Scientific Reports |language=en |volume=7 |issue=1 |pages=1572 |doi=10.1038/s41598-017-01837-7 |issn=2045-2322}}</ref>
* [[Niger Congo|Niger-Congo]] and [[Nilo-Saharan languages|Nilo-Saharan]] speakers split around 28,000 years ago.<ref name=":5" />
* [[Austronesian languages|Austronesian-speaking]] [[Malagasy people]] in [[Madagascar]] have received significant [[East Asian people|East/Southeast Asian]] admixture, less among some groups of coastal Southern, Eastern and the Horn of Africa. The estimated date of geneflow is 2,200 years ago.<ref>{{Cite journal |last1=Kusuma |first1=Pradiptajati |last2=Brucato |first2=Nicolas |last3=Cox |first3=Murray P. |last4=Pierron |first4=Denis |last5=Razafindrazaka |first5=Harilanto |last6=Adelaar |first6=Alexander |last7=Sudoyo |first7=Herawati |last8=Letellier |first8=Thierry |last9=Ricaut |first9=François-Xavier |date=2016-05-18 |title=Contrasting Linguistic and Genetic Origins of the Asian Source Populations of Malagasy |journal=Scientific Reports |volume=6 |pages=26066 |doi=10.1038/srep26066 |issn=2045-2322 |pmc=4870696 |pmid=27188237|bibcode=2016NatSR...626066K }}</ref><ref>{{cite journal |vauthors=Cruciani F, Santolamazza P, Shen P, Macaulay V, Moral P, Olckers A, Modiano D, Holmes S, Destro-Bisol G, Coia V, Wallace DC, Oefner PJ, Torroni A, Cavalli-Sforza LL, Scozzari R, Underhill PA |display-authors=6 |title=A back migration from Asia to sub-Saharan Africa is supported by high-resolution analysis of human Y-chromosome haplotypes |journal=American Journal of Human Genetics |volume=70 |issue=5 |pages=1197–1214 |date=May 2002 |pmid=11910562 |pmc=447595 |doi=10.1086/340257}}</ref><ref>{{cite journal |vauthors=Pakstis AJ, Speed WC, Soundararajan U, Rajeevan H, Kidd JR, Li H, Kidd KK |title=Population relationships based on 170 ancestry SNPs from the combined Kidd and Seldin panels |journal=Scientific Reports |volume=9 |issue=1 |pages=18874 |date=December 2019 |pmid=31827153 |pmc=6906462 |doi=10.1038/s41598-019-55175-x |bibcode=2019NatSR...918874P}}</ref><ref name="Fan 82" /><ref name="Identifying and Interpreting Appare">{{cite journal |vauthors=Chen L, Wolf AB, Fu W, Li L, Akey JM |title=Identifying and Interpreting Apparent Neanderthal Ancestry in African Individuals |journal=Cell |volume=180 |issue=4 |pages=677–687.e16 |date=February 2020 |pmid=32004458 |doi=10.1016/j.cell.2020.01.012 |s2cid=210955842}}</ref><ref name=":2">{{Cite journal |last1=van de Loosdrecht |first1=Marieke |last2=Bouzouggar |first2=Abdeljalil |last3=Humphrey |first3=Louise |last4=Posth |first4=Cosimo |last5=Barton |first5=Nick |last6=Aximu-Petri |first6=Ayinuer |last7=Nickel |first7=Birgit |last8=Nagel |first8=Sarah |last9=Talbi |first9=El Hassan |last10=El Hajraoui |first10=Mohammed Abdeljalil |last11=Amzazi |first11=Saaïd |last12=Hublin |first12=Jean-Jacques |last13=Pääbo |first13=Svante |last14=Schiffels |first14=Stephan |last15=Meyer |first15=Matthias |date=2018-05-04 |title=Pleistocene North African genomes link Near Eastern and sub-Saharan African human populations |url=https://www.science.org/doi/10.1126/science.aar8380 |journal=Science |language=en |volume=360 |issue=6388 |pages=548–552 |doi=10.1126/science.aar8380 |pmid=29545507 |bibcode=2018Sci...360..548V |s2cid=206666517 |issn=0036-8075}}</ref>
[[File:Overall genetic position of worldwide populations.jpg|thumb|Geographic location of the samples analyzed in this study (A). PCA of the Khoe-San individuals, Eurasians, West and East Africans before (unmasked, B) and after (masked, C) applying the local ancestry pipeline (146,696 independent SNPs).]]
===Indigenous Africans===
The indigenous population of Africa consists of [[Niger–Congo languages|Niger–Congo speakers]], [[Nilo-Saharan languages|Nilo-Saharan speakers]], the divergent and diverse [[Khoisan languages|Khoisan grouping]], as well as of several unclassified or [[Language isolate|isolated]] ethnolinguistic groupings (see [[:Category:Unclassified languages of Africa|unclassified languages of Africa]]). The origin of the [[Afroasiatic languages]] remains disputed, with some proposing a [[Middle Eastern]] origin, while others support an African origin with varying degrees of Eurasian and African components.<ref>{{Cite journal |last=Baker |first=Jennifer L. |last2=Rotimi |first2=Charles N. |last3=Shriner |first3=Daniel |date=2017-05-08 |title=Human ancestry correlates with language and reveals that race is not an objective genomic classifier |url=https://www.nature.com/articles/s41598-017-01837-7 |journal=Scientific Reports |language=en |volume=7 |issue=1 |pages=1572 |doi=10.1038/s41598-017-01837-7 |issn=2045-2322}}</ref> The [[Austronesian languages]] originated in southern East Asia, and later expanded outgoing from the [[Philippines]].
[[File:Procrustes-transformed PCA plot of genetic variation of worldwide populations.png|thumb|PCA plot of genetic variation of worldwide populations. ('''A''') Geographic coordinates of 53 populations. ('''B''') Procrustes-transformed PCA plot of genetic variation.<ref>{{cite journal |vauthors=Wang C, Zöllner S, Rosenberg NA |title=A quantitative comparison of the similarity between genes and geography in worldwide human populations |journal=PLOS Genetics |volume=8 |issue=8 |pages=e1002886 |date=August 2012 |pmid=22927824 |pmc=3426559 |doi=10.1371/journal.pgen.1002886}}</ref>]]
The [[Niger–Congo languages]] probably originated in or near the area where these languages were spoken prior to [[Bantu expansion]] (i.e. [[West Africa]] or [[Central Africa]]). Its expansion may have been associated with the expansion of agriculture, during in the [[Neolithic|African Neolithic]] period, following the [[African humid period#End|desiccation of the Sahara]] in c. 3500 BCE. [[Proto-Niger-Congo]] may have originated about 10,000 years before present in the "[[Green Sahara]]" of Africa (roughly the [[Sahel]] and southern [[Sahara]]), and that its dispersal can be correlated with the spread of the [[bow and arrow]] by migrating [[hunter-gatherer]]s, which later developed agriculture.<ref>{{Cite journal |vauthors=Manning K, Timpson A |date=2014-10-01 |title=The demographic response to Holocene climate change in the Sahara |journal=Quaternary Science Reviews |language=en |volume=101 |pages=28–35 |doi=10.1016/j.quascirev.2014.07.003 |bibcode=2014QSRv..101...28M |s2cid=54923700 |issn=0277-3791}}</ref><ref name="BlenchAfricanPast">{{Cite book |vauthors=Blench R |title=Archaeology, language, and the African past |publisher=AltaMira Press |year=2006 |isbn=9780759104655}}</ref><ref>{{cite conference |vauthors=Blench R |url=https://llacan.cnrs.fr/nigercongo2/discussions/Can_we_visit_the_graves_of_the_first_Niger.pdf |title=Can we visit the graves of the first Niger-Congo speakers? |conference=2nd International Congress: Towards Proto-Niger-Congo: Comparison and Reconstruction |location=Paris |date=September 2016}}</ref>
Although the validity of the [[Nilo-Saharan languages|Nilo-Saharan family]] remains controversial, the region between [[Chad]], [[Sudan]], and the [[Central African Republic]] is seen as a likely candidate for its homeland prior to its dispersal around 10,000–8,000 BCE.<ref>{{Cite book |vauthors=Vossen R, Dimmendaal GJ |url=https://books.google.com/books?id=wcjXDwAAQBAJ&pg=PA380 |title=The Oxford Handbook of African Languages |date=2020-03-13 |publisher=Oxford University Press |isbn=978-0-19-960989-5 |language=en}}</ref>
The Southern African hunter-gatherers (Khoisan) are suggested to represent the autochthonous hunter-gatherer population of southern Africa, prior to the expansion of Bantu-speakers from Western/Central Africa and East African pastoralists. Khoisan show evidence for Bantu-related admixture, ranging from nearly ~0% to up to ~87.1%.<ref>{{cite journal |vauthors=Vicente M, Jakobsson M, Ebbesen P, Schlebusch CM |title=Genetic Affinities among Southern Africa Hunter-Gatherers and the Impact of Admixing Farmer and Herder Populations |journal=Molecular Biology and Evolution |volume=36 |issue=9 |pages=1849–1861 |date=September 2019 |pmid=31288264 |pmc=6735883 |doi=10.1093/molbev/msz089}}</ref>
===Out-of-Africa event===
[[File:Human migration routes following Out-of-Africa.png|thumb|Human migration routes following Out-of-Africa.]]
[[File:Migraciones humanas en haplogrupos de ADN-Y.PNG|thumb|Most modern Africans display a high level of genetic homogeneity, but contributions from Eurasian populations are substantial, mostly concentrated in the Northeastern part of Africa and Madagascar.]]
The model proposes a "single origin" of ''[[Homo sapiens]]'' in Africa the taxonomic sense. Recent genetic and archeologic data suggests that Homo sapiens-subgroups originated in multiple regions of Africa, not confined to a single region of origin. The ''H. sapiens'' ancestral to proper Eurasians most likely left [[Northeastern Africa]] between 50,000 and 100,000 years ago.<ref name=":2" /> The "recent African origin" model proposes that all modern non-African populations descend from one or several diverse waves of ''H. sapiens'' that left Africa ~70,000 years ago, with some proposing an earlier date.<ref>{{cite journal |vauthors=López S, van Dorp L, Hellenthal G |title=Human Dispersal Out of Africa: A Lasting Debate |journal=Evolutionary Bioinformatics Online |volume=11 |issue=Suppl 2 |pages=57–68 |date=2016-04-21 |pmid=27127403 |pmc=4844272 |doi=10.4137/EBO.S33489}}</ref><ref>{{cite journal |vauthors=Scerri EM, Thomas MG, Manica A, Gunz P, Stock JT, Stringer C, Grove M, Groucutt HS, Timmermann A, Rightmire GP, d'Errico F, Tryon CA, Drake NA, Brooks AS, Dennell RW, Durbin R, Henn BM, Lee-Thorp J, deMenocal P, Petraglia MD, Thompson JC, Scally A, Chikhi L |display-authors=6 |title=Did Our Species Evolve in Subdivided Populations across Africa, and Why Does It Matter? |language=English |journal=Trends in Ecology & Evolution |volume=33 |issue=8 |pages=582–594 |date=August 2018 |pmid=30007846 |pmc=6092560 |doi=10.1016/j.tree.2018.05.005}}</ref><ref>{{Cite web |title=One Species, Many Origins |url=https://www.shh.mpg.de/1474609/pan-african-origins |access-date=2022-01-11 |website=www.shh.mpg.de |language=en}}</ref>
According to a 2020 study by Durvasula et al., there are indications that 2% to 19% (≃6.6 to 7.0%) of the DNA of West African populations may have come from an unknown archaic hominin which split from the ancestor of humans and Neanderthals between 360 kya to 1.02 mya. However, the study also suggests that at least part of this archaic admixture is also present in Eurasians/non-Africans, and that the admixture event or events range from 0 to 124 ka B.P, which includes the period before the Out-of-Africa migration and prior to the African/Eurasian split (thus affecting in part the common ancestors of both Africans and Eurasians/non-Africans).<ref>{{cite journal |vauthors=Durvasula A, Sankararaman S |date=February 2020 |title=Recovering signals of ghost archaic introgression in African populations |journal=Science Advances |volume=6 |issue=7 |pages=eaax5097 |bibcode=2020SciA....6.5097D |doi=10.1126/sciadv.aax5097 |pmc=7015685 |pmid=32095519 |doi-access=free}} "Non-African populations (Han Chinese in Beijing and Utah residents with northern and western European ancestry) also show analogous patterns in the CSFS, suggesting that a component of archaic ancestry was shared before the split of African and non-African populations...One interpretation of the recent time of introgression that we document is that archaic forms persisted in Africa until fairly recently. Alternately, the archaic population could have introgressed earlier into a modern human population, which then subsequently interbred with the ancestors of the populations that we have analyzed here. The models that we have explored here are not mutually exclusive, and it is plausible that the history of African populations includes genetic contributions from multiple divergent populations, as evidenced by the large effective population size associated with the introgressing archaic population...Given the uncertainty in our estimates of the time of introgression, we wondered whether jointly analyzing the CSFS from both the CEU (Utah residents with Northern and Western European ancestry) and YRI genomes could provide additional resolution. Under model C, we simulated introgression before and after the split between African and non-African populations and observed qualitative differences between the two models in the high-frequency–derived allele bins of the CSFS in African and non-African populations (fig. S40). Using ABC to jointly fit the high-frequency–derived allele bins of the CSFS in CEU and YRI (defined as greater than 50% frequency), we find that the lower limit on the 95% credible interval of the introgression time is older than the simulated split between CEU and YRI (2800 versus 2155 generations B.P.), indicating that at least part of the archaic lineages seen in the YRI are also shared with the CEU..."</ref><ref>''[https://www.science.org/doi/10.1126/sciadv.aax5097] {{Webarchive|url=https://web.archive.org/web/20201207165127/https://advances.sciencemag.org/content/advances/suppl/2020/02/10/6.7.eaax5097.DC1/aax5097_SM.pdf|date=7 December 2020}} Supplementary Materials for ''Recovering signals of ghost archaic introgression in African populations", section "S8.2" "We simulated data using the same priors in Section S5.2, but computed the spectrum for both YRI [West African Yoruba] and CEU [a population of European origin] . We found that the best fitting parameters were an archaic split time of 27,000 generations ago (95% HPD: 26,000-28,000), admixture fraction of 0.09 (95% HPD: 0.04-0.17), admixture time of 3,000 generations ago (95% HPD: 2,800-3,400), and an effective population size of 19,700 individuals (95% HPD: 19,300-20,200). We find that the lower bound of the admixture time is further back than the simulated split between CEU and YRI (2155 generations ago), providing some evidence in favor of a pre-Out-of-Africa event. This model suggests that many populations outside of Africa should also contain haplotypes from this introgression event, though detection is difficult because many methods use unadmixed outgroups to detect introgressed haplotypes [Browning et al., 2018, Skov et al., 2018, Durvasula and Sankararaman, 2019] (5, 53, 22). It is also possible that some of these haplotypes were lost during the Out-of-Africa bottleneck."</ref><ref>{{cite journal |vauthors=Durvasula A, Sankararaman S |date=February 2020 |title=Recovering signals of ghost archaic introgression in African populations |journal=Science Advances |volume=6 |issue=7 |pages=eaax5097 |bibcode=2020SciA....6.5097D |doi=10.1126/sciadv.aax5097 |pmc=7015685 |pmid=32095519}}</ref> Another academic paper from 2020 (Chen et al.) found that Africans have higher [[Neanderthal]] ancestry than previously thought. 2,504 African samples from all over Africa were analyzed and tested on Neanderthal ancestry. All African samples showed evidence for minor Neanderthal ancestry, but always at lower levels than observed in Eurasians.<ref name="Identifying and Interpreting Appare2">{{cite journal |vauthors=Chen L, Wolf AB, Fu W, Li L, Akey JM |date=February 2020 |title=Identifying and Interpreting Apparent Neanderthal Ancestry in African Individuals |journal=Cell |volume=180 |issue=4 |pages=677–687.e16 |doi=10.1016/j.cell.2020.01.012 |pmid=32004458 |s2cid=210955842}}</ref>
===Geneflow between Eurasian and African populations===
{{See also|Eurasian backflow}}
[[File:Pre-Neolithic and Neolithic migration events in Africa (excluding Basal-West-Eurasian geneflow during the Paleolithic).jpg|thumb|Pre-Neolithic and Neolithic migration events in Africa.<ref>{{Cite journal |last=Vicente |first=Mário |last2=Schlebusch |first2=Carina M |date=2020-06-01 |title=African population history: an ancient DNA perspective |url=https://www.sciencedirect.com/science/article/pii/S0959437X20300599 |journal=Current Opinion in Genetics & Development |series=Genetics of Human Origin |language=en |volume=62 |pages=8–15 |doi=10.1016/j.gde.2020.05.008 |issn=0959-437X}}</ref>]]
Significant Eurasian admixture is found in [[Northern Africa]], and among specific ethnic groups of the [[Horn of Africa]], as well as among the [[Malagasy people]] of [[Madagascar]]. Various genome studies found evidence for multiple [[Prehistory|prehistoric]] back-migrations from various [[Eurasia]]n populations and subsequent admixture with native groups.<ref>{{cite journal |vauthors=Busby GB, Band G, Si Le Q, Jallow M, Bougama E, Mangano VD, Amenga-Etego LN, Enimil A, Apinjoh T, Ndila CM, Manjurano A, Nyirongo V, Doumba O, Rockett KA, Kwiatkowski DP, Spencer CC |display-authors=6 |title=Admixture into and within sub-Saharan Africa |journal=eLife |volume=5 |date=June 2016 |pmid=27324836 |pmc=4915815 |doi=10.7554/eLife.15266}}</ref> West-Eurasian geneflow arrived to Northern Africa during the Paleolithic (30,000 to 15,000 years ago), followed by other pre-Neolithic and Neolithic migration events. Genetic data on the [[Taforalt]] samples "demonstrated that Northern Africa received significant amounts of gene-flow from Eurasia predating the Holocene and development of farming practices". Medieval geneflow events, such as the [[Arab expansion]] also left traces in various African populations.<ref>{{Cite journal |last=van de Loosdrecht |first=Marieke |last2=Bouzouggar |first2=Abdeljalil |last3=Humphrey |first3=Louise |last4=Posth |first4=Cosimo |last5=Barton |first5=Nick |last6=Aximu-Petri |first6=Ayinuer |last7=Nickel |first7=Birgit |last8=Nagel |first8=Sarah |last9=Talbi |first9=El Hassan |last10=El Hajraoui |first10=Mohammed Abdeljalil |last11=Amzazi |first11=Saaïd |last12=Hublin |first12=Jean-Jacques |last13=Pääbo |first13=Svante |last14=Schiffels |first14=Stephan |last15=Meyer |first15=Matthias |date=2018-05-04 |title=Pleistocene North African genomes link Near Eastern and sub-Saharan African human populations |url=https://www.science.org/doi/10.1126/science.aar8380 |journal=Science |language=en |volume=360 |issue=6388 |pages=548–552 |doi=10.1126/science.aar8380 |issn=0036-8075}}</ref><ref>{{Cite journal |last=Vicente |first=Mário |last2=Schlebusch |first2=Carina M |date=2020-06-01 |title=African population history: an ancient DNA perspective |url=https://www.sciencedirect.com/science/article/pii/S0959437X20300599 |journal=Current Opinion in Genetics & Development |series=Genetics of Human Origin |language=en |volume=62 |pages=8–15 |doi=10.1016/j.gde.2020.05.008 |issn=0959-437X}}</ref><ref>{{Cite journal |last=Serra-Vidal |first=Gerard |last2=Lucas-Sanchez |first2=Marcel |last3=Fadhlaoui-Zid |first3=Karima |last4=Bekada |first4=Asmahan |last5=Zalloua |first5=Pierre |last6=Comas |first6=David |date=2019-11-18 |title=Heterogeneity in Palaeolithic Population Continuity and Neolithic Expansion in North Africa |url=https://www.sciencedirect.com/science/article/pii/S0960982219312412 |journal=Current Biology |language=en |volume=29 |issue=22 |pages=3953–3959.e4 |doi=10.1016/j.cub.2019.09.050 |issn=0960-9822}}</ref> A study (Pickrell et al. 2014) indicated that Western Eurasian ancestry eventually arrived through [[Northeast Africa]] (particularly the Horn of Africa) to [[Southern Africa]].<ref name=":3">{{cite journal |display-authors=6 |vauthors=Pickrell JK, Patterson N, Loh PR, Lipson M, Berger B, Stoneking M, Pakendorf B, Reich D |date=February 2014 |title=Ancient west Eurasian ancestry in southern and eastern Africa |journal=Proceedings of the National Academy of Sciences of the United States of America |volume=111 |issue=7 |pages=2632–2637 |arxiv=1307.8014 |bibcode=2014PNAS..111.2632P |doi=10.1073/pnas.1313787111 |pmc=3932865 |pmid=24550290 |doi-access=free}}</ref>
Another study (Ramsay ''et al.'' 2018) also found evidence for significant Western Eurasian admixture in various parts of Africa, from both ancient and more recent migrations, being highest among populations from [[North Africa|Northern Africa]], and some groups of the [[Horn of Africa]]:<ref name="Ramsay">{{cite journal |vauthors=Choudhury A, Aron S, Sengupta D, Hazelhurst S, Ramsay M |title=African genetic diversity provides novel insights into evolutionary history and local adaptations |journal=Human Molecular Genetics |volume=27 |issue=R2 |pages=R209–R218 |date=August 2018 |pmid=29741686 |pmc=6061870 |doi=10.1093/hmg/ddy161}}</ref>
<blockquote>
In addition to the intrinsic diversity within the continent due to population structure and isolation, migration of Eurasian populations into Africa has emerged as a critical contributor to the genetic diversity. These migrations involved the influx of different Eurasian populations at different times and to different parts of Africa. Comprehensive characterization of the details of these migrations through genetic studies on existing populations could help to explain the strong genetic differences between some geographically neighbouring populations.
This distinctive Eurasian admixture appears to have occurred over at least three time periods with ancient admixture in central west Africa (e.g. Yoruba from Nigeria) occurring between ~7.5 and 10.5 kya, older admixture in east Africa (e.g. Ethiopia) occurring between ~2.4 and 3.2 kya and more recent admixture between ~0.15 and 1.5 kya in some east African (e.g. Kenyan) populations.
Subsequent studies based on LD decay and haplotype sharing in an extensive set of African and Eurasian populations confirmed the presence of Eurasian signatures in west, east and southern Africans. In the west, in addition to Niger-Congo speakers from The Gambia and Mali, the Mossi from Burkina Faso showed the oldest Eurasian admixture event ~7 kya. In the east, these analyses inferred Eurasian admixture within the last 4000 years in Kenya.</blockquote>
[[File:Expansion of Afroasiatic.svg|thumb|Proposed migration and expansion routes of the [[Afroasiatic languages]] according to the indigenous African origin model.]]
There is no definitive agreement on when or where the [[Afroasiatic homeland|original homeland]] of the [[Afroasiatic languages|Afroasiatic language family]] existed. Some have suggested that they were spread by people with largely West-Eurasian ancestry during the [[Neolithic Revolution]], towards Northern Africa and the Horn of Africa, outgoing from the [[Middle East]], specifically from the [[Levant]]. This hypothesis does not account for the domestication of plants [[Endemism|endemic]] to the Horn of Africa such as [[teff]], [[Ensete ventricosum|ensete]], and [[Guizotia abyssinica|niger seed]], nor does it account for the lack of evidence of intrusive agricultural populations or the cultivation of [[wheat]], [[barley]], or [[sorghum]] in that region prior to 3000 B.C.<ref>{{Cite book |last1=Clark |first1=JD |title=From Hunters to Farmers: The Causes and Consequences of Food Production in Africa |last2=Brandt |first2=SA |publisher=University of California Press |year=1984 |isbn=978-0520045743 |pages=180 |language=English}}</ref><ref>{{cite journal |vauthors=Diamond J, Bellwood P |date=April 2003 |title=Farmers and their languages: the first expansions |journal=Science |volume=300 |issue=5619 |pages=597–603 |bibcode=2003Sci...300..597D |doi=10.1126/science.1078208 |pmid=12714734 |s2cid=13350469}}</ref> Others argue that the first speakers of Afroasiatic ([[Proto-Afroasiatic]]) were based in Northeast Africa because that region includes the majority of the diversity of the Afroasiatic language family and has very diverse groups in close geographic proximity, sometimes considered a telltale sign for a linguistic geographic origin.<ref>{{Cite book |last=Campbell |first=Lyle |title=Historical Linguistics, Fourth Edition |publisher=The MIT Press |year=2021 |isbn=978-0262542180 |pages=399–400 |language=English}}</ref> A subset of the Proto-Afroasiatic population would have migrated to the [[Levant]] during the late [[Paleolithic]], merging with local West-Eurasians and resulting in a population which would later give rise to [[Natufian culture]], associated with the early development of [[agriculture]] and early Afroasiatic languages, or specifically pre-[[Proto-Semitic language|proto-Semitic]].<ref>{{Cite book |last1=Jarvie |title=Transition to Modernity: Essays on Power, Wealth and Belief |last2=Hall |publisher=Cambridge University Press |year=2005 |isbn=9780521022279 |pages=27 |language=English}}</ref><ref>{{Cite book |last=Shirai |first=Noriyuki |url=http://worldcat.org/oclc/852516752 |title=The archaeology of the first farmer-herders in Egypt: new insights into the Fayum Epipalaeolithic and Neolithic |date=2010 |publisher=Leiden University Press |isbn=978-90-485-1269-0 |oclc=852516752}}</ref>{{Page needed|date=March 2022}}<ref name="Early back-to-Africa migration into">{{cite journal |vauthors=Hodgson JA, Mulligan CJ, Al-Meeri A, Raaum RL |date=June 2014 |title=Early back-to-Africa migration into the Horn of Africa |journal=PLOS Genetics |volume=10 |issue=6 |pages=e1004393 |doi=10.1371/journal.pgen.1004393 |pmc=4055572 |pmid=24921250}}</ref><ref>{{Cite book |last=Blench |first=Roger |title=Archaeology, Language, and the African Past |publisher=AltaMira Press |year=2006 |isbn=978-0759104662 |pages=150–163 |language=English}}</ref><ref name=":4">{{Cite journal |last=Ehret |first=Christopher |date=1979 |title=On the Antiquity of Agriculture in Ethiopia |url=https://www.jstor.org/stable/181512 |journal=The Journal of African History |volume=20 |issue=2 |pages=161–177 |doi=10.1017/S002185370001700X |jstor=181512 |s2cid=162986221 |via=JSTOR}}</ref><ref>{{Cite book |last=Nöth |first=Winfried |title=Origins of Semiosis: Sign Evolution in Nature and Culture |publisher=De Gruyter Mouton |year=2011 |isbn=9781134816231 |pages=293 |language=English}}</ref>
[[File:E of Y-DNA migrations.png|thumb|Proposed migration routes of paternal lineage E.]]
While many studies conducted on Horn of Africa populations estimate a West-Eurasian admixture event around 3,000 years ago,<ref name=":0" /><ref name="Ramsay" /><ref name=":3" /><ref>{{Cite journal |last1=Molinaro |first1=Ludovica |last2=Montinaro |first2=Francesco |last3=Yelmen |first3=Burak |last4=Marnetto |first4=Davide |last5=Behar |first5=Doron M. |last6=Kivisild |first6=Toomas |last7=Pagani |first7=Luca |date=2019-12-11 |title=West Asian sources of the Eurasian component in Ethiopians: a reassessment |journal=Scientific Reports |language=en |volume=9 |issue=1 |pages=18811 |doi=10.1038/s41598-019-55344-y |pmid=31827175 |pmc=6906521 |bibcode=2019NatSR...918811M |issn=2045-2322}}</ref> a 2014 genome study by Hodgson et al. found a distinct West-Eurasian ancestral component among studied Afroasiatic-speaking groups in the Horn of Africa (and to a lesser extent in North Africa and [[Western Asia|West Asia]]), most prevalent among the [[Somalis|Somali]]. This ancestral component — dubbed "Ethio-Somali" — would have diverged from other non-African ancestries around 23,000 years ago and migrated back to Africa prior to developing agriculture, merging with the local indigenous lineages of the Horn of Africa. The authors propose that "Ethio-Somali" may have been a substantial ancestral component of the Proto-Afroasiatic-speaking population. A subsequent [[Mitochondrial DNA|mtDNA]] (Gandini et al. 2016) analysis has produced additional evidence in support of a pre-agricultural back-migration from West-Eurasia into the [[Horn of Africa]] with an estimated date of arrival into the Horn of Africa in the early [[Holocene]], possibly as a result of [[obsidian]] exchange networks across the Red Sea.<ref>{{Cite journal |last1=Gandini |first1=Francesca |last2=Achilli |first2=Alessandro |last3=Pala |first3=Maria |last4=Bodner |first4=Martin |last5=Brandini |first5=Stefania |last6=Huber |first6=Gabriela |last7=Egyed |first7=Balazs |last8=Ferretti |first8=Luca |last9=Gómez-Carballa |first9=Alberto |last10=Salas |first10=Antonio |last11=Scozzari |first11=Rosaria |date=2016-05-05 |title=Mapping human dispersals into the Horn of Africa from Arabian Ice Age refugia using mitogenomes |journal=Scientific Reports |language=en |volume=6 |issue=1 |pages=25472 |doi=10.1038/srep25472 |pmid=27146119 |pmc=4857117 |bibcode=2016NatSR...625472G |issn=2045-2322}}</ref> Hodgson et al. also confirmed the existence of an ancestral component indigenous to the Horn of Africa – "Ethiopic" (Hodgson et al.) or "Omotic" (Pagani et al.) – which is most prevalent among speakers of the [[Omotic languages|Omotic]] branch of Afroasiatic in southwestern Ethiopia.<ref name="Early back-to-Africa migration into" /><ref name=":0" /> This lineage is associated with that of a 4,500 year-old fossil (Mota) found in a cave in southwestern Ethiopia, which has high genetic affinity to modern [[Ethiopians|Ethiopian groups]], especially the [[Endogamy|endogamous]] blacksmith [[caste]] of the Omotic [[Ari people|Aari people]]. Like Mota, Aari blacksmiths do not show evidence for admixture with West-Eurasians, demonstrating a degree of population continuity in this region for at least 4,500 years. In a comparative analysis of Mota’s genome referencing modern populations, Gallego et al. (2016) concluded that the divergence of Omotic from other Afroasiatic languages may have resulted from the relative isolation of its speakers from external groups.<ref>{{Cite journal |date=2016-02-19 |title=Erratum for the Report "Ancient Ethiopian genome reveals extensive Eurasian admixture in Eastern Africa" (previously titled "Ancient Ethiopian genome reveals extensive Eurasian admixture throughout the African continent") by M. Gallego Llorente, E. R. Jones, A. Eriksson, V. Siska, K. W. Arthur, J. W. Arthur, M. C. Curtis, J. T. Stock, M. Coltorti, P. Pieruccini, S. Stretton, F. Brock, T. Higham, Y. Park, M. Hofreiter, D. G. Bradley, J. Bhak, R. Pinhasi, A. Manica |url=https://pubmed.ncbi.nlm.nih.gov/26912899/ |journal=Science |volume=351 |issue=6275 |pages=aaf3945 |doi=10.1126/science.aaf3945 |issn=1095-9203 |pmid=26912899}}</ref> In an analysis of 68 Ethiopian ethnic groups, also referencing Mota, Lopez et al. (2021) revealed that groups belonging to the Omotic, Cushitic, and Semitic branches of Afro-Asiatic show high genetic similarity to each other on average. The data also support widespread recent intermixing among ethnic groups.<ref>{{Cite journal |last1=López |first1=Saioa |last2=Tarekegn |first2=Ayele |last3=Band |first3=Gavin |last4=van Dorp |first4=Lucy |last5=Bird |first5=Nancy |last6=Morris |first6=Sam |last7=Oljira |first7=Tamiru |last8=Mekonnen |first8=Ephrem |last9=Bekele |first9=Endashaw |last10=Blench |first10=Roger |last11=Thomas |first11=Mark G. |date=2021-06-11 |title=Evidence of the interplay of genetics and culture in Ethiopia |journal=Nature Communications |language=en |volume=12 |issue=1 |pages=3581 |doi=10.1038/s41467-021-23712-w |pmid=34117245 |pmc=8196081 |bibcode=2021NatCo..12.3581L |issn=2041-1723}}</ref> Linguist [[Roger Blench]] proposed southwestern Ethiopia as the most likely homeland of Afroasiatic, due in part to the high internal diversification of the Omotic branch spoken in that region.<ref name="BlenchAfricanPast" /> In addition, [[Human Y-chromosome DNA haplogroup|Y-haplogroup]] sub-lineage [[Haplogroup E-M215 (Y-DNA)|E-M215]] (also known as "E1b1b) and its derivative E-M35 are quite common among Afroasiatic speakers and southwestern Ethiopia is a plausible source of these haplogroups.<ref name=":1">{{Cite journal |last=Shriner |first=Daniel |date=2018 |title=Re-analysis of Whole Genome Sequence Data From 279 Ancient Eurasians Reveals Substantial Ancestral Heterogeneity |journal=Frontiers in Genetics |volume=9 |page=268 |doi=10.3389/fgene.2018.00268 |issn=1664-8021 |pmc=6062619 |pmid=30079081 |quote=and a sub-Saharan African component in Natufians that localizes to present-day southern Ethiopia. |doi-access=free}}</ref> The linguistic group and carriers of this lineage have a high probability to have arisen and dispersed together from Northeast Africa in the [[Mesolithic]], plausibly having already developed [[subsistence pattern]]s of [[pastoralism]] and intensive plant usage and collection.<ref>{{Cite journal |last1=Underhill |first1=P. A. |last2=Passarino |first2=G. |last3=Lin |first3=A. A. |last4=Shen |first4=P. |last5=Mirazón Lahr |first5=M. |last6=Foley |first6=R. A. |last7=Oefner |first7=P. J. |last8=Cavalli-Sforza |first8=L. L. |date=January 2001 |title=The phylogeography of Y chromosome binary haplotypes and the origins of modern human populations |journal=Annals of Human Genetics |volume=65 |issue=Pt 1 |pages=43–62 |doi=10.1046/j.1469-1809.2001.6510043.x |issn=0003-4800 |pmid=11415522 |s2cid=9441236}}</ref><ref>{{Cite journal |last=Ibrahim |first=Muntaser E. |date=2021-04-26 |title=Genetic diversity of the Sudanese: insights on origin and implications for health |journal=Human Molecular Genetics |volume=30 |issue=R1 |pages=R37–R41 |doi=10.1093/hmg/ddab028 |issn=1460-2083 |pmc=8223596 |pmid=33864377}}</ref><ref>{{Cite journal |last1=Ehret |first1=Christopher |last2=Keita |first2=S. O. Y. |last3=Newman |first3=Paul |date=2004-12-03 |title=The origins of Afroasiatic |journal=Science |volume=306 |issue=5702 |pages=1680; author reply 1680 |doi=10.1126/science.306.5702.1680c |issn=1095-9203 |pmid=15576591 |s2cid=8057990}}</ref><ref>{{Cite journal |last=Ehret |first=Christopher |date=1979 |title=On the Antiquity of Agriculture in Ethiopia |journal=The Journal of African History |volume=20 |issue=2 |pages=161–177 |doi=10.1017/S002185370001700X |jstor=181512 |s2cid=162986221 |issn=0021-8537}}</ref> According to historian and linguist [[Christopher Ehret]], the form of intensive plant collection practiced by the Proto-Afroasiatic population in Northeast Africa may have been a precursor to the agricultural practices that would later independently develop in the [[Fertile Crescent]] and the Horn of Africa.<ref name=":4"/><ref>{{Citation |last1=Bultosa |first1=G. |title=TEFF |date=2004-01-01 |url=https://www.sciencedirect.com/science/article/pii/B0127654909001725 |encyclopedia=Encyclopedia of Grain Science |pages=281–290 |editor-last=Wrigley |editor-first=Colin |place=Oxford |publisher=Elsevier |language=en |doi=10.1016/b0-12-765490-9/00172-5 |isbn=978-0-12-765490-4 |access-date=2022-03-29 |last2=Taylor |first2=J. R. N.}}</ref><ref>{{Cite journal |last1=Schlebusch |first1=Carina M. |last2=Jakobsson |first2=Mattias |date=2018-08-31 |title=Tales of Human Migration, Admixture, and Selection in Africa |journal=Annual Review of Genomics and Human Genetics |volume=19 |pages=405–428 |doi=10.1146/annurev-genom-083117-021759 |issn=1545-293X |pmid=29727585 |s2cid=19155657}}</ref> A 2018 re-analysis of autosomal DNA using modern populations as a reference found that the ancient Natufian samples of the Levant harbored 6.8% Omotic-related ancestry.<ref name=":1"/> A 2015 study by Dobon et al. identified an autosomal ancestral component that is commonly found among modern Afroasiatic-speaking populations (as well as [[Nubians]]) in Northeast Africa. This component, which peaks among [[Copts]] in [[Sudan]] but is not found in [[Egyptians]] or [[Qataris]], appears alongside a component that defines [[Nilo-Saharan languages|Nilo-Saharan]] speakers of southwestern Sudan and [[South Sudan]].<ref>{{cite journal |display-authors=6 |vauthors=Dobon B, Hassan HY, Laayouni H, Luisi P, Ricaño-Ponce I, Zhernakova A, Wijmenga C, Tahir H, Comas D, Netea MG, Bertranpetit J |date=May 2015 |title=The genetics of East African populations: a Nilo-Saharan component in the African genetic landscape |journal=Scientific Reports |volume=5 |pages=9996 |bibcode=2015NatSR...5E9996D |doi=10.1038/srep09996 |pmc=4446898 |pmid=26017457}}</ref> A 2017 paper by Arauna et al. analyzing existing genetic data obtained from [[North Africa|Northern African]] populations, such as [[Berbers]], described them as a mosaic of Middle Eastern, European, and [[Sub-Saharan Africa]]n-related ancestries.<ref>{{Citation |vauthors=Arauna LR, Comas D |title=Genetic Heterogeneity between Berbers and Arabs |date=2017 |work=eLS |pages=1–7 |publisher=John Wiley & Sons, Ltd |language=en |doi=10.1002/9780470015902.a0027485 |isbn=978-0-470-01590-2}}</ref>
[[File:Austronesia with hypothetical greatest expansion extent (Blench, 2009) 01.png|thumb|Austronesian expansion, outgoing from [[Taiwan]] and the northern [[Philippines]].]]
Specific East Asian-related ancestry is found among the [[Malagasy languages|Malagasy speakers]] of [[Madagascar]] at a medium frequency. The presence of this East Asian-related ancestry is mostly linked to the [[Austronesian peoples]] expansion from [[Southeast Asia]].<ref>{{Cite journal |vauthors=Dewar RE, Wright HT |date=1993-12-01|title=The culture history of Madagascar |journal=Journal of World Prehistory |language=en |volume=7 |issue=4 |pages=417–466 |doi=10.1007/BF00997802 |hdl=2027.42/45256 |s2cid=21753825 |hdl-access=free}}</ref><ref>{{cite journal | vauthors = Burney DA, Burney LP, Godfrey LR, Jungers WL, Goodman SM, Wright HT, Jull AJ | title = A chronology for late prehistoric Madagascar | journal = Journal of Human Evolution | volume = 47 | issue = 1–2 | pages = 25–63 | date = 2004-07-01 | pmid = 15288523 | doi = 10.1016/j.jhevol.2004.05.005 }}</ref><ref>{{cite journal | vauthors = Pierron D, Razafindrazaka H, Pagani L, Ricaut FX, Antao T, Capredon M, Sambo C, Radimilahy C, Rakotoarisoa JA, Blench RM, Letellier T, Kivisild T | display-authors = 6 | title = Genome-wide evidence of Austronesian-Bantu admixture and cultural reversion in a hunter-gatherer group of Madagascar | journal = Proceedings of the National Academy of Sciences of the United States of America | volume = 111 | issue = 3 | pages = 936–941 | date = January 2014 | pmid = 24395773 | pmc = 3903192 | doi = 10.1073/pnas.1321860111 | doi-access = free | bibcode = 2014PNAS..111..936P }}</ref><ref name="Kusuma_2016">{{cite journal | vauthors = Kusuma P, Brucato N, Cox MP, Pierron D, Razafindrazaka H, Adelaar A, Sudoyo H, Letellier T, Ricaut FX | display-authors = 6 | title = Contrasting Linguistic and Genetic Origins of the Asian Source Populations of Malagasy | journal = Scientific Reports | volume = 6 | issue = 1 | pages = 26066 | date = May 2016 | pmid = 27188237 | pmc = 4870696 | doi = 10.1038/srep26066 | bibcode = 2016NatSR...626066K }}</ref> The peoples of [[Borneo]] were identified to resemble the East Asian voyagers, who arrived on Madagascar. East Asian ancestry among Malagasy people was estimated at a mean average of 33%, but as high as ~75% among some Highlander groups and upper caste groups.<ref name="Heiske"/><ref>{{Cite web |last=Nicolas |first=Brucato |display-authors=etal |date=4 February 2019 |title=Evidence of Austronesian Genetic Lineages in East Africa and South Arabia: Complex Dispersal from Madagascar and Southeast Asia |url=https://academic.oup.com/gbe/article/11/3/748/5306180 |website=Genome Biology and Evolution, Volume 11, Issue 3}}</ref><ref name="Kusuma_2016" />
A 2020 study by Chen et al. analyzed 2,504 African samples from all over Africa, and found archaic Neanderthal ancestry, among all tested African samples at low frequency. They also identified a European-related (West-Eurasian) ancestry segment, which seems to largely correspond with the detected Neanderthal ancestry components. European-related admixture among Africans was estimated to be between ~0% to up to ~30%, with a peak among Northern Africans.<ref name="Identifying and Interpreting Appare"/>
According to the authors:<ref name="Identifying and Interpreting Appare"/>
{{Blockquote|text=These data are consistent with the hypothesis that back-migration contributed to the signal of Neanderthal ancestry in Africans. Furthermore, the data indicates that this back-migration came after the split of Europeans and East Asians, from a population related to the European lineage."}}
A 2021 paper by Hollfelder et al., concluded that West African [[Yoruba people]], which were previously used as "unadmixed reference population" for indigenous Africans, harbor minor levels of Neanderthal ancestry, which can be largely associated with back-migration of an "Ancestral European-like" source population.<ref name="The deep population history in Afri"/>
[[File:Map_of_African_admixture_in_European_populations.png|thumb|Map of [[Sub-Saharan African]] and specific [[North African]] admixture in European populations]]
Multiple studies found also evidence for geneflow of indigenous African ancestry towards Eurasia, specifically Europe and the Middle East. The analysis of 40 different West-Eurasian populations found African admixture at a frequency of 0% to up to ~15%.<ref>{{Cite journal |last1=Moorjani |first1=Priya |last2=Patterson |first2=Nick |last3=Hirschhorn |first3=Joel N. |last4=Keinan |first4=Alon |last5=Hao |first5=Li |last6=Atzmon |first6=Gil |last7=Burns |first7=Edward |last8=Ostrer |first8=Harry |last9=Price |first9=Alkes L. |last10=Reich |first10=David |date=April 2011 |title=The history of African gene flow into Southern Europeans, Levantines, and Jews |journal=PLOS Genetics |volume=7 |issue=4 |pages=e1001373 |doi=10.1371/journal.pgen.1001373 |issn=1553-7404 |pmc=3080861 |pmid=21533020}}</ref><ref>{{Cite journal |last1=Botigué |first1=Laura R. |last2=Henn |first2=Brenna M. |last3=Gravel |first3=Simon |last4=Maples |first4=Brian K. |last5=Gignoux |first5=Christopher R. |last6=Corona |first6=Erik |last7=Atzmon |first7=Gil |last8=Burns |first8=Edward |last9=Ostrer |first9=Harry |last10=Flores |first10=Carlos |last11=Bertranpetit |first11=Jaume |date=2013-07-16 |title=Gene flow from North Africa contributes to differential human genetic diversity in southern Europe |journal=Proceedings of the National Academy of Sciences of the United States of America |volume=110 |issue=29 |pages=11791–11796 |doi=10.1073/pnas.1306223110 |issn=0027-8424 |pmc=3718088 |pmid=23733930 |bibcode=2013PNAS..11011791B |doi-access=free }}</ref><ref>{{Cite journal |last1=Auton |first1=Adam |last2=Bryc |first2=Katarzyna |last3=Boyko |first3=Adam R. |last4=Lohmueller |first4=Kirk E. |last5=Novembre |first5=John |last6=Reynolds |first6=Andy |last7=Indap |first7=Amit |last8=Wright |first8=Mark H. |last9=Degenhardt |first9=Jeremiah D. |last10=Gutenkunst |first10=Ryan N. |last11=King |first11=Karen S. |date=May 2009 |title=Global distribution of genomic diversity underscores rich complex history of continental human populations |journal=Genome Research |volume=19 |issue=5 |pages=795–803 |doi=10.1101/gr.088898.108 |issn=1088-9051 |pmc=2675968 |pmid=19218534}}</ref><ref>{{Cite journal |last1=Reich |first1=David |last2=Price |first2=Alkes L. |last3=Patterson |first3=Nick |date=May 2008 |title=Principal component analysis of genetic data |url=https://www.nature.com/articles/ng0508-491 |journal=Nature Genetics |language=en |volume=40 |issue=5 |pages=491–492 |doi=10.1038/ng0508-491 |pmid=18443580 |s2cid=34837532 |issn=1546-1718}}</ref>
Evidence for minor geneflow from West or West-Central Africa into the [[Iberian Peninsula]] is supported by the presence of an African-specific mitochondrial haplogroup among one of four 4,000 year old samples.<ref>{{Cite journal |last1=González-Fortes |first1=G. |last2=Tassi |first2=F. |last3=Trucchi |first3=E. |last4=Henneberger |first4=K. |last5=Paijmans |first5=J. L. A. |last6=Díez-del-Molino |first6=D. |last7=Schroeder |first7=H. |last8=Susca |first8=R. R. |last9=Barroso-Ruíz |first9=C. |last10=Bermudez |first10=F. J. |last11=Barroso-Medina |first11=C. |last12=Bettencourt |first12=A. M. S. |last13=Sampaio |first13=H. A. |last14=Grandal-d'Anglade |first14=A. |last15=Salas |first15=A. |date=2019-01-30 |title=A western route of prehistoric human migration from Africa into the Iberian Peninsula |journal=Proceedings of the Royal Society B: Biological Sciences |volume=286 |issue=1895 |pages=20182288 |doi=10.1098/rspb.2018.2288 |issn=0962-8452 |pmc=6364581 |pmid=30963949}}</ref>
==Regional genomic overview==
===North Africa===
{{Main|Genetic history of North Africa}}
{{Further|DNA history of Egypt|Genetic studies on Moroccans}}
====Archaic Human DNA====
{{Further|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.<ref name="Bergström">{{cite journal | vauthors = Bergström A, Stringer C, Hajdinjak M, Scerri EM, Skoglund P | title = Origins of modern human ancestry | journal = Nature | volume = 590 | issue = 7845 | pages = 229–237 | date = February 2021 | pmid = 33568824 | doi = 10.1038/s41586-021-03244-5 | s2cid = 231883210 | bibcode = 2021Natur.590..229B }}</ref>
====Ancient DNA====
An early medieval sample from the 5th to 6th century CE found in northern France (Angers) was found to be similar to present-day North Africans.<ref>{{Cite journal |last=Alves |first=Isabel |last2=Giemza |first2=Joanna |last3=Blum |first3=Michael |last4=Bernhardsson |first4=Carolina |last5=Chatel |first5=Stéphanie |last6=Karakachoff |first6=Matilde |last7=Pierre |first7=Aude Saint |last8=Herzig |first8=Anthony F. |last9=Olaso |first9=Robert |last10=Monteil |first10=Martial |last11=Gallien |first11=Véronique |last12=Cabot |first12=Elodie |last13=Svensson |first13=Emma |last14=Bacq-Daian |first14=Delphine |last15=Baron |first15=Estelle |date=2022-02-04 |title=Genetic population structure across Brittany and the downstream Loire basin provides new insights on the demographic history of Western Europe |url=https://www.biorxiv.org/content/10.1101/2022.02.03.478491v1 |language=en |pages=2022.02.03.478491 |doi=10.1101/2022.02.03.478491v1.full}}</ref>
=====Egypt=====
[[Tomb of Two Brothers#The tomb owners|Khnum-aa]], [[Tomb of Two Brothers|Khnum-Nakht]], [[Tomb of Two Brothers#Coffins of Nakht-Ankh|Nakht-Ankh]] and JK2911 carried maternal [[Haplogroup M (mtDNA)#Haplogroup M1|haplogroup M1a1]].<ref name="Gad" /><ref name=":0">{{Cite journal |last1=Schuenemann |first1=Verena J. |last2=Peltzer |first2=Alexander |last3=Welte |first3=Beatrix |last4=van Pelt |first4=W. Paul |last5=Molak |first5=Martyna |last6=Wang |first6=Chuan-Chao |last7=Furtwängler |first7=Anja |last8=Urban |first8=Christian |last9=Reiter |first9=Ella |last10=Nieselt |first10=Kay |last11=Teßmann |first11=Barbara |date=2017-05-30 |title=Ancient Egyptian mummy genomes suggest an increase of Sub-Saharan African ancestry in post-Roman periods |journal=Nature Communications |volume=8 |issue=1 |page=15694 |doi=10.1038/ncomms15694 |pmid=28556824 |issn=2041-1723 |doi-access=free |bibcode=2017NatCo...815694S |pmc=5459999}}</ref>
[[Djehutynakht (10A)]] carried maternal haplogroup [[Haplogroup U (mtDNA)#Haplogroup U5|U5b2b5]].<ref name="Loreille 135">{{Cite journal |last1=Loreille |first1=Odile |last2=Ratnayake |first2=Shashikala |last3=Bazinet |first3=Adam L. |last4=Stockwell |first4=Timothy B. |last5=Sommer |first5=Daniel D. |last6=Rohland |first6=Nadin |last7=Mallick |first7=Swapan |last8=Johnson |first8=Philip L. F. |last9=Skoglund |first9=Pontus |last10=Onorato |first10=Anthony J. |last11=Bergman |first11=Nicholas H. |date=March 2018 |title=Biological Sexing of a 4000-Year-Old Egyptian Mummy Head to Assess the Potential of Nuclear DNA Recovery from the Most Damaged and Limited Forensic Specimens |journal=Genes |volume=9 |issue=3 |pages=135 |doi=10.3390/genes9030135 |pmid=29494531 |pmc=5867856 |doi-access=free}}</ref> JK2888 carried maternal haplogroup [[Haplogroup U (mtDNA)|U6a2]].<ref name=":0" />
[[Thuya]], [[Tiye]], Tutankhamen's mother, and Tutankhamen carried the maternal [[Haplogroup K (mtDNA)|haplogroup K]].<ref name="Gad" />
JK2134 carried maternal haplogroup [[Haplogroup J (mtDNA)|J1d]]<ref name=":0" /> and JK2887 carried maternal haplogroup [[Haplogroup J (mtDNA)|J2a1a1]].<ref name=":0" />
[[Amenhotep III]], [[Akhenaten]], and [[Tutankhamen]] carried the paternal [[haplogroup R1b]].<ref name="Gad">{{cite journal |vauthors=Gad YZ, Hassan NA, Mousa DM, Fouad FA, El-Sayed SG, Abdelazeem MA, Mahdy SM, Othman HY, Ibrahim DW, Khairat R, Ismail S |display-authors=6 |title=Insights from ancient DNA analysis of Egyptian human mummies: clues to disease and kinship |journal=Human Molecular Genetics |volume=30 |issue=R1 |pages=R24–R28 |date=April 2021 |pmid=33059357 |doi=10.1093/hmg/ddaa223 |doi-access=free}}</ref>
[[Ramesses III]] and "Unknown Man E", possibly [[Pentawere]], carried paternal [[haplogroup E1b1a]].<ref name="Gad" /><ref name="Hawass">{{cite journal |vauthors=Hawass Z, Ismail S, Selim A, Saleem SN, Fathalla D, Wasef S, Gad AZ, Saad R, Fares S, Amer H, Gostner P, Gad YZ, Pusch CM, Zink AR |display-authors=6 |title=Revisiting the harem conspiracy and death of Ramesses III: anthropological, forensic, radiological, and genetic study |journal=BMJ |volume=345 |pages=e8268 |date=December 2012 |pmid=23247979 |doi=10.1136/bmj.e8268 |hdl-access=free |s2cid=206896841 |hdl=10072/62081}}</ref><ref name="Gourdine">{{cite journal | vauthors = Gourdine JP, Keita S, Gourdine JL, Anselin A |title=Ancient Egyptian Genomes from northern Egypt: Further discussion |url=https://www.researchgate.net/publication/327065612 |journal=Nature Communications}}</ref>
JK2134 and JK2911 carried paternal haplogroup [[Haplogroup J (Y-DNA)|J]].<ref name=":0" />
[[Takabuti]] carried maternal haplogroup [[Haplogroup H (mtDNA)|H4a1]]<ref>{{Cite web |url=https://www.manchester.ac.uk/discover/news/shocking-truth-behind-takabutis-death-revealed/ |title=Shocking truth behind Takabuti's death revealed |language=en |access-date=2020-01-29}}</ref> and YM:KMM A 63 carried maternal haplogroup [[Haplogroup HV. (mtDNA)|HV]]<ref name=":10">{{Cite journal |last1=Oras |first1=Ester |last2=Anderson |first2=Jaanika |last3=Tõrv |first3=Mari |last4=Vahur |first4=Signe |last5=Rammo |first5=Riina |last6=Remmer |first6=Sünne |last7=Mölder |first7=Maarja |last8=Malve |first8=Martin |last9=Saag |first9=Lehti |last10=Saage |first10=Ragnar |last11=Teearu-Ojakäär |first11=Anu |date=2020-01-16 |title=Multidisciplinary investigation of two Egyptian child mummies curated at the University of Tartu Art Museum, Estonia (Late/Graeco-Roman Periods) |journal=PLOS ONE |language=en |volume=15 |issue=1 |pages=e0227446 |doi=10.1371/journal.pone.0227446 |pmid=31945091 |pmc=6964855 |bibcode=2020PLoSO..1527446O |issn=1932-6203 |doi-access=free}}</ref>
OM:KMM A 64 carried maternal haplogroup [[Haplogroup T (mtDNA)|T2c1a]].<ref name=":10" />
JK2888 carried paternal haplogroup [[Haplogroup E-M215 (Y-DNA)|E1b1b1a1b2]].<ref name=":0" />
=====Libya=====
At Takarkori rockshelter, in [[Libya]], two naturally [[mummified]] women, dated to the [[Pastoral Period#Middle Pastoral Period 2|Middle Pastoral Period]] (7000 BP), carried [[Basal (phylogenetics)|basal]] maternal [[Haplogroup N (mtDNA)|haplogroup N]].<ref name="Vai">{{cite journal | vauthors = Vai S, Sarno S, Lari M, Luiselli D, Manzi G, Gallinaro M, Mataich S, Hübner A, Modi A, Pilli E, Tafuri MA, Caramelli D, di Lernia S | display-authors = 6 | title = Ancestral mitochondrial N lineage from the Neolithic 'green' Sahara | journal = Scientific Reports | volume = 9 | issue = 1 | pages = 3530 | date = March 2019 | pmid = 30837540 | pmc = 6401177 | doi = 10.1038/s41598-019-39802-1 | bibcode = 2019NatSR...9.3530V }}</ref>
=====Morocco=====
A 2018 study by Van de Loorsdrecht et al. found that of seven samples of [[Taforalts]] of [[Morocco]], [[radiocarbon dated]] to between 15,100 cal BP and 13,900 cal BP, six were found to carry maternal haplogroup [[Haplogroup U (mtDNA)|U6a]], and one was found to carry maternal haplogroup [[Haplogroup M (mtDNA)|M1b]]. Six of six males were found to carry paternal haplogroup [[Haplogroup E-M215 (Y-DNA)|E1b1b]]. They were found to harbor 63.5% [[Natufian]]-related ancestry and 36.5% [[Sub-Saharan African]]-related ancestry. The Sub-Saharan component is most strongly drawn out by modern West African groups such as the [[Yoruba people|Yoruba]] and the [[Mende people|Mende]]. The samples also contain an additional affinity to South, Central, and East African outgroups that cannot be explained by any known ancient or modern populations.<ref name="Van De Loosdrecht">{{cite journal | vauthors = van de Loosdrecht M, Bouzouggar A, Humphrey L, Posth C, Barton N, Aximu-Petri A, Nickel B, Nagel S, Talbi EH, El Hajraoui MA, Amzazi S, Hublin JJ, Pääbo S, Schiffels S, Meyer M, Haak W, Jeong C, Krause J | display-authors = 6 | title = Pleistocene North African genomes link Near Eastern and sub-Saharan African human populations | journal = Science | volume = 360 | issue = 6388 | pages = 548–552 | date = May 2018 | pmid = 29545507 | doi = 10.1126/science.aar8380 | s2cid = 206666517 | doi-access = free | bibcode = 2018Sci...360..548V }}</ref> When projected onto a principal component analysis graph of African and west Eurasian populations, the Taforalt individuals form a distinct cluster in an intermediate position between present-day North Africans [e.g., [[Berbers|Amazighes]] (Berbers), Mozabites, and Saharawis] and East Africans (e.g., [[Afar people|Afars]], Oromos, and Somalis).<ref name="Van De Loosdrecht"/> Another study (Jeong 2020) comparing the Taforalt people of the [[Iberomaurusian]] culture to modern populations found that the Taforalt Sub-Saharan African genetic component may be best represented by modern West Africans (e.g. Yoruba).<ref name="Jeong" />
====Y-Chromosomal DNA====
{{Further|Y-DNA haplogroups in populations of North Africa}}
====Mitochondrial DNA====
Mitochondrial haplogroups L3, M, and N are found among [[Sudan#Ethnic groups|Sudanese peoples]] (e.g., [[Beja people|Beja]], [[Nilotic peoples|Nilotics]], [[Nuba]], [[Nubians]]), who have no known interaction (e.g., history of migration/admixture) with Europeans or Asians; rather than having developed in a post-Out-of-Africa migration context, mitochondrial macrohaplogroup L3/M/N and its subsequent development into distinct mitochondrial haplogroups (e.g., [[Haplogroup L3]], [[Haplogroup M (mtDNA)|Haplogroup M]], [[Haplogroup N (mtDNA)|Haplogroup N]]) may have occurred in [[East Africa]] at a time that considerably predates the Out-of-Africa migration event of 50,000 BP.<ref name="Osman">{{cite journal |vauthors=Osman MM, Hassan HY, Elnour MA, Makkan H, Gebremeskel EI, Gais T, Koko ME, Soodyall H, Ibrahim ME | display-authors = 6 |title=Mitochondrial HVRI and whole mitogenome sequence variations portray similar scenarios on the genetic structure and ancestry of northeast Africans |url=http://81.95.108.158/return-files/Mitochondrial%20HVRI.pdf |journal=Meta Gene}}</ref>
====Autosomal DNA====
====Medical DNA====
=====Lactase Persistence=====
[[Neolithic]] [[agriculturalists]], who may have resided in [[Northeast Africa]] and the [[Near East]], may have been the source population for [[lactase persistence]] variants, including –13910*T, and may have been subsequently supplanted by later migrations of peoples.<ref name="Priehodová">{{cite journal |vauthors=Priehodová E, Austerlitz F, Čížková M, Nováčková J, Ricaut FX, Hofmanová Z, Schlebusch CM, Černý V | display-authors = 6 | title = Sahelian pastoralism from the perspective of variants associated with lactase persistence | journal = American Journal of Physical Anthropology | volume = 173 | issue = 3 | pages = 423–436 | date = November 2020 | pmid = 32812238 | doi = 10.1002/ajpa.24116 | s2cid = 221179656 | url = https://hal.archives-ouvertes.fr/hal-02919786/file/ajpa_ms_final.pdf }}</ref> The [[Sub-Saharan]] [[West African]] Fulani, the [[North African]] [[Tuareg]], and [[Early European Farmers|European agriculturalists]], who are descendants of these Neolithic agriculturalists, share the lactase persistence variant –13910*T.<ref name="Priehodová" /> While shared by Fulani and Tuareg herders, compared to the Tuareg variant, the Fulani variant of –13910*T has undergone a longer period of haplotype differentiation.<ref name="Priehodová" /> The [[Fulani]] lactase persistence variant –13910*T may have spread, along with cattle [[pastoralism]], between 9686 BP and 7534 BP, possibly around 8500 BP; corroborating this timeframe for the Fulani, by at least 7500 BP, there is evidence of herders engaging in the act of [[milking]] in the Central [[Sahara]].<ref name="Priehodová" />
===West Africa===
====Archaic Human DNA====
{{Further|Interbreeding between archaic and modern humans#Archaic African hominins}}
Archaic traits found in human fossils of [[West Africa]] (e.g., [[Iwo Eleru skull|Iho Eleru fossils]], which dates to 13,000 BP) and [[Central Africa]] (e.g., [[Ishango]] fossils, which dates between 25,000 BP and 20,000 BP) may have developed as a result of admixture between archaic humans and modern humans or may be evidence of late-persisting [[Early modern human#Early Homo sapiens|early modern humans]].<ref name="Bergström"/> 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.<ref name="Bergström" />
====Ancient DNA====
As of 2017, human ancient DNA has not been found in the region of [[Western Africa]].<ref name="Scerri">{{cite web | vauthors = Scerri E |title= The Stone Age Archaeology of West Africa |url=https://ora.ox.ac.uk/objects/uuid:1442d1c1-7f76-454d-97db-8fd52a4c6c1e/download_file?file_format=pdf&safe_filename=The%2BStone%2BAge%2BArchaeology%2Bof%2BWest%2BAfrica.pdf&type_of_work=Book+section |website=Oxford University Research Archive |publisher=Oxford University Press}}</ref> As of 2020, human ancient DNA has not been forthcoming in the region of Western Africa.<ref name="Jeong">{{cite book | vauthors = Jeong C |chapter=Current Trends in Ancient DNA Study |chapter-url=https://link.springer.com/referenceworkentry/10.1007%2F978-981-15-1614-6_10-1 |year=2020 |pages=1–16 |title=The Handbook of Mummy Studies|publisher=Springer |doi=10.1007/978-981-15-1614-6_10-1 |isbn=978-981-15-1614-6 |s2cid=226555687 }}</ref>
In 4000 BP, there may have been a population that traversed from [[Africa]] (e.g., [[West Africa]] or West-[[Central Africa]]), through the [[Strait of Gibraltar]], into the [[Iberian peninsula]], where admixing between Africans and Iberians (e.g., of northern [[Portugal]], of southern [[Spain]]) occurred.<ref name="RSP">{{cite journal | vauthors = González-Fortes G, Tassi F, Trucchi E, Henneberger K, Paijmans JL, Díez-Del-Molino D, Schroeder H, Susca RR, Barroso-Ruíz C, Bermudez FJ, Barroso-Medina C, Bettencourt AM, Sampaio HA, Grandal-d'Anglade A, Salas A, de Lombera-Hermida A, Fabregas Valcarce R, Vaquero M, Alonso S, Lozano M, Rodríguez-Alvarez XP, Fernández-Rodríguez C, Manica A, Hofreiter M, Barbujani G | display-authors = 6 | title = A western route of prehistoric human migration from Africa into the Iberian Peninsula | journal = Proceedings. Biological Sciences | volume = 286 | issue = 1895 | pages = 20182288 | date = January 2019 | pmid = 30963949 | pmc = 6364581 | doi = 10.1098/rspb.2018.2288 }}</ref>
In [[Granada]], a Muslim ([[Moors|Moor]]) of the [[Cordoba Caliphate]],<ref name="Olalde">{{cite journal | vauthors = Olalde I, Mallick S, Patterson N, Rohland N, Villalba-Mouco V, Silva M, Dulias K, Edwards CJ, Gandini F, Pala M, Soares P, Ferrando-Bernal M, Adamski N, Broomandkhoshbacht N, Cheronet O, Culleton BJ, Fernandes D, Lawson AM, Mah M, Oppenheimer J, Stewardson K, Zhang Z, Jiménez Arenas JM, Toro Moyano IJ, Salazar-García DC, Castanyer P, Santos M, Tremoleda J, Lozano M, García Borja P, Fernández-Eraso J, Mujika-Alustiza JA, Barroso C, Bermúdez FJ, Viguera Mínguez E, Burch J, Coromina N, Vivó D, Cebrià A, Fullola JM, García-Puchol O, Morales JI, Oms FX, Majó T, Vergès JM, Díaz-Carvajal A, Ollich-Castanyer I, López-Cachero FJ, Silva AM, Alonso-Fernández C, Delibes de Castro G, Jiménez Echevarría J, Moreno-Márquez A, Pascual Berlanga G, Ramos-García P, Ramos-Muñoz J, Vijande Vila E, Aguilella Arzo G, Esparza Arroyo Á, Lillios KT, Mack J, Velasco-Vázquez J, Waterman A, Benítez de Lugo Enrich L, Benito Sánchez M, Agustí B, Codina F, de Prado G, Estalrrich A, Fernández Flores Á, Finlayson C, Finlayson G, Finlayson S, Giles-Guzmán F, Rosas A, Barciela González V, García Atiénzar G, Hernández Pérez MS, Llanos A, Carrión Marco Y, Collado Beneyto I, López-Serrano D, Sanz Tormo M, Valera AC, Blasco C, Liesau C, Ríos P, Daura J, de Pedro Michó MJ, Diez-Castillo AA, Flores Fernández R, Francès Farré J, Garrido-Pena R, Gonçalves VS, Guerra-Doce E, Herrero-Corral AM, Juan-Cabanilles J, López-Reyes D, McClure SB, Merino Pérez M, Oliver Foix A, Sanz Borràs M, Sousa AC, Vidal Encinas JM, Kennett DJ, Richards MB, Werner Alt K, Haak W, Pinhasi R, Lalueza-Fox C, Reich D | display-authors = 6 | title = The genomic history of the Iberian Peninsula over the past 8000 years | journal = Science | volume = 363 | issue = 6432 | pages = 1230–1234 | date = March 2019 | pmid = 30872528 | pmc = 6436108 | doi = 10.1126/science.aav4040 | bibcode = 2019Sci...363.1230O | hdl = 10261/207967 }}</ref> who was of haplogroups [[Haplogroup E-M2|E1b1a1]] and [[Haplogroup H (mtDNA)#H1|H1+16189]],<ref name="Olalde II">{{cite journal | vauthors = Olalde I |title=Ancient individuals from the Iberian Peninsula included in this study |url=https://science.sciencemag.org/highwire/filestream/724016/field_highwire_adjunct_files/2/aav4040_TablesS1-S5.xlsx |journal=Science}}</ref><ref name="Olalde III">{{cite journal | vauthors = Olalde I, Mallick S, Patterson N, Rohland N, Villalba-Mouco V, Silva M, Dulias K, Edwards CJ, Gandini F, Pala M, Soares P, Ferrando-Bernal M, Adamski N, Broomandkhoshbacht N, Cheronet O, Culleton BJ, Fernandes D, Lawson AM, Mah M, Oppenheimer J, Stewardson K, Zhang Z, Jiménez Arenas JM, Toro Moyano IJ, Salazar-García DC, Castanyer P, Santos M, Tremoleda J, Lozano M, García Borja P, Fernández-Eraso J, Mujika-Alustiza JA, Barroso C, Bermúdez FJ, Viguera Mínguez E, Burch J, Coromina N, Vivó D, Cebrià A, Fullola JM, García-Puchol O, Morales JI, Oms FX, Majó T, Vergès JM, Díaz-Carvajal A, Ollich-Castanyer I, López-Cachero FJ, Silva AM, Alonso-Fernández C, Delibes de Castro G, Jiménez Echevarría J, Moreno-Márquez A, Pascual Berlanga G, Ramos-García P, Ramos-Muñoz J, Vijande Vila E, Aguilella Arzo G, Esparza Arroyo Á, Lillios KT, Mack J, Velasco-Vázquez J, Waterman A, Benítez de Lugo Enrich L, Benito Sánchez M, Agustí B, Codina F, de Prado G, Estalrrich A, Fernández Flores Á, Finlayson C, Finlayson G, Finlayson S, Giles-Guzmán F, Rosas A, Barciela González V, García Atiénzar G, Hernández Pérez MS, Llanos A, Carrión Marco Y, Collado Beneyto I, López-Serrano D, Sanz Tormo M, Valera AC, Blasco C, Liesau C, Ríos P, Daura J, de Pedro Michó MJ, Diez-Castillo AA, Flores Fernández R, Francès Farré J, Garrido-Pena R, Gonçalves VS, Guerra-Doce E, Herrero-Corral AM, Juan-Cabanilles J, López-Reyes D, McClure SB, Merino Pérez M, Oliver Foix A, Sanz Borràs M, Sousa AC, Vidal Encinas JM, Kennett DJ, Richards MB, Werner Alt K, Haak W, Pinhasi R, Lalueza-Fox C, Reich D | display-authors = 6 | title = The genomic history of the Iberian Peninsula over the past 8000 years | journal = Science | volume = 363 | issue = 6432 | pages = 1230–1234 | date = March 2019 | pmid = 30872528 | pmc = 6436108 | doi = 10.1126/science.aav4040 | bibcode = 2019Sci...363.1230O }}</ref> as well as estimated to date between 900 CE and 1000 CE, and a [[Morisco]],<ref name="Olalde" /> who was of [[Haplogroup L2 (mtDNA)#Haplogroup L2e|haplogroup L2e1]],<ref name="Olalde II" /><ref name="Olalde III" /> as well as estimated to date between 1500 CE and 1600 CE, were both found to be of Sub-Saharan West African (i.e., [[The Gambia#Ethnic groups|Gambian]]), [[Iberian Peninsula|Iberian]], and North African descent.<ref name="Olalde" />
====Y-Chromosomal DNA====
{{Further|Y-DNA haplogroups in populations of Sub-Saharan Africa}}
As a result of haplogroup D0, a basal branch of haplogroup DE, being found in three [[Nigerian]] men, it may be the case that [[haplogroup DE]], as well as its sublineages D0 and E, originated in [[Africa]].<ref name="Haber">{{cite journal | vauthors = Haber M, Jones AL, Connell BA, Arciero E, Yang H, Thomas MG, Xue Y, Tyler-Smith C | display-authors = 6 | title = A Rare Deep-Rooting D0 African Y-Chromosomal Haplogroup and Its Implications for the Expansion of Modern Humans Out of Africa | journal = Genetics | volume = 212 | issue = 4 | pages = 1421–1428 | date = August 2019 | pmid = 31196864 | pmc = 6707464 | doi = 10.1534/genetics.119.302368 }}</ref>
As of 19,000 years ago, Africans, bearing [[Haplogroup E-V38|haplogroup E1b1a-V38]], likely traversed across the [[Sahara]], from [[East Africa|east]] to [[West Africa|west]].<ref name="Shriner">{{cite journal | vauthors = Shriner D, Rotimi CN | title = Whole-Genome-Sequence-Based Haplotypes Reveal Single Origin of the Sickle Allele during the Holocene Wet Phase | journal = American Journal of Human Genetics | volume = 102 | issue = 4 | pages = 547–556 | date = April 2018 | pmid = 29526279 | pmc = 5985360 | doi = 10.1016/j.ajhg.2018.02.003 }}</ref> [[Haplogroup E-M2|E1b1a1-M2]] likely originated in [[West Africa]] or [[Central Africa]].<ref name="Trombetta">{{cite journal | vauthors = Trombetta B, D'Atanasio E, Massaia A, Ippoliti M, Coppa A, Candilio F, Coia V, Russo G, Dugoujon JM, Moral P, Akar N, Sellitto D, Valesini G, Novelletto A, Scozzari R, Cruciani F | display-authors = 6 | title = Phylogeographic Refinement and Large Scale Genotyping of Human Y Chromosome Haplogroup E Provide New Insights into the Dispersal of Early Pastoralists in the African Continent | journal = Genome Biology and Evolution | volume = 7 | issue = 7 | pages = 1940–1950 | date = June 2015 | pmid = 26108492 | pmc = 4524485 | doi = 10.1093/gbe/evv118 }}</ref>
====Mitochondrial DNA====
{{Further|Haplogroup L2 (mtDNA)}}
Around 18,000 BP, [[Mende people]], along with [[The Gambia#Ethnic groups|Gambian peoples]], grew in population size.<ref name="Miller">{{cite journal | vauthors = Miller EF, Manica A, Amos W | title = Global demographic history of human populations inferred from whole mitochondrial genomes | journal = Royal Society Open Science | volume = 5 | issue = 8 | pages = 180543 | date = August 2018 | pmid = 30225046 | pmc = 6124094 | doi = 10.1098/rsos.180543 | bibcode = 2018RSOS....580543M }}</ref>
In 15,000 BP, [[Niger-Congo]] speakers may have migrated from the [[Sahelian]] region of West Africa, along the [[Senegal River]], and introduced [[Haplogroup L2 (mtDNA)#Haplogroup L2a1|L2a1]] into [[North Africa]], resulting in modern [[Demographics of Mauritania#Ethnic groups|Mauritanian peoples]] and [[Berbers]] of [[Tunisia]] inheriting it.<ref name="Frigi">{{cite journal | vauthors = Frigi S, Cherni L, Fadhlaoui-Zid K, Benammar-Elgaaied A | title = Ancient local evolution of African mtDNA haplogroups in Tunisian Berber populations | journal = Human Biology | volume = 82 | issue = 4 | pages = 367–384 | date = August 2010 | pmid = 21082907 | doi = 10.3378/027.082.0402 | s2cid = 27594333 }}</ref>
Between 11,000 BP and 10,000 BP, [[Yoruba people]] and [[Esan people]] grew in population size.<ref name="Miller" />
Up to 11,000 years ago, Sub-Saharan West Africans, bearing [[Macro-haplogroup L (mtDNA)|macrohaplogroup L]] (e.g., [[Haplogroup L1 (mtDNA)#L1b|L1b1a11]], L1b1a6a, L1b1a8, L1b1a9a1, [[Haplogroup L2 (mtDNA)#Haplogroup L2a1|L2a1k]], [[Haplogroup L3 (mtDNA)#Subclade distribution|L3d1b1a]]), may have migrated through [[North Africa]] and into [[Europe]], mostly into [[southern Europe]] (e.g., [[Iberia]]).<ref name="Soares">{{cite journal | vauthors = Podgorná E, Soares P, Pereira L, Cerný V | title = The genetic impact of the lake chad basin population in North Africa as documented by mitochondrial diversity and internal variation of the L3e5 haplogroup | journal = Annals of Human Genetics | volume = 77 | issue = 6 | pages = 513–523 | date = November 2013 | pmid = 25069842 | doi = 10.1111/ahg.12040 | s2cid = 24672148 }}</ref>
====Autosomal DNA====
Between 2000 BP and 1500 BP, [[Nilo-Saharan]]-speakers may have migrated across the [[Sahel]], from [[East Africa]] into [[West Africa]], and admixed with [[Niger-Congo]]-speaking [[Berom people]].<ref name="Choudhury II">{{cite journal | vauthors = Choudhury A, Aron S, Botigué LR, Sengupta D, Botha G, Bensellak T, Wells G, Kumuthini J, Shriner D, Fakim YJ, Ghoorah AW, Dareng E, Odia T, Falola O, Adebiyi E, Hazelhurst S, Mazandu G, Nyangiri OA, Mbiyavanga M, Benkahla A, Kassim SK, Mulder N, Adebamowo SN, Chimusa ER, Muzny D, Metcalf G, Gibbs RA, Rotimi C, Ramsay M, Adeyemo AA, Lombard Z, Hanchard NA | display-authors = 6 | title = High-depth African genomes inform human migration and health | journal = Nature | volume = 586 | issue = 7831 | pages = 741–748 | date = October 2020 | pmid = 33116287 | doi = 10.1038/s41586-020-2859-7 | pmc = 7759466 | bibcode = 2020Natur.586..741C }}</ref>
A 2019 study on [[Fulani people]] by Fan et al., found that the Fulani show genetic affinity to isolated [[Afroasiatic languages|Afroasiatic-speaking]] groups in [[East Africa|Eastern Africa]], specifically [[Omotic languages|Omotic-speakers]] such as the [[Aari people]]. While the Fulani have nearly exclusive indigenous African ancestry (defined by West and East African ancestry), they also show traces of West-Eurasian-like admixture, supporting an ancestral homeland somewhere in North or Eastern Africa, and westwards expansion during the Neolithic, possibly caused by the arrival and expansion of West-Eurasian-related groups.<ref>{{Cite journal |last1=Fan |first1=Shaohua |last2=Kelly |first2=Derek E. |last3=Beltrame |first3=Marcia H. |last4=Hansen |first4=Matthew E. B. |last5=Mallick |first5=Swapan |last6=Ranciaro |first6=Alessia |last7=Hirbo |first7=Jibril |last8=Thompson |first8=Simon |last9=Beggs |first9=William |last10=Nyambo |first10=Thomas |last11=Omar |first11=Sabah A. |date=2019-04-26 |title=African evolutionary history inferred from whole genome sequence data of 44 indigenous African populations |url=https://doi.org/10.1186/s13059-019-1679-2 |journal=Genome Biology |volume=20 |issue=1 |pages=82 |doi=10.1186/s13059-019-1679-2 |issn=1474-760X |pmc=6485071 |pmid=31023338}}</ref>
====Medical DNA====
=====Sickle Cell=====
Amid the Green Sahara, the mutation for [[sickle cell]] originated in the [[Sahara]]<ref name="Shriner" /> or in the [[Northwestern Congolian lowland forests|northwest forest]] region of western Central Africa (e.g., Cameroon)<ref name="Shriner" /><ref name="Esoh">{{cite journal | vauthors = Esoh K, Wonkam A | title = Evolutionary history of sickle-cell mutation: implications for global genetic medicine | journal = Human Molecular Genetics | volume = 30 | issue = R1 | pages = R119–R128 | date = April 2021 | pmid = 33461216 | pmc = 8117455 | doi = 10.1093/hmg/ddab004 }}</ref> by at least 7,300 years ago,<ref name="Shriner" /><ref name="Esoh" /> though possibly as early as 22,000 years ago.<ref name="Laval">{{cite journal | vauthors = Laval G, Peyrégne S, Zidane N, Harmant C, Renaud F, Patin E, Prugnolle F, Quintana-Murci L | display-authors = 6 | title = Recent Adaptive Acquisition by African Rainforest Hunter-Gatherers of the Late Pleistocene Sickle-Cell Mutation Suggests Past Differences in Malaria Exposure | journal = American Journal of Human Genetics | volume = 104 | issue = 3 | pages = 553–561 | date = March 2019 | pmid = 30827499 | pmc = 6407493 | doi = 10.1016/j.ajhg.2019.02.007 }}</ref><ref name="Esoh" /> The ancestral sickle cell haplotype to modern haplotypes (e.g., [[Cameroon]]/[[Central African Republic]] and [[Benin]]/[[Senegal]] haplotypes) may have first arose in the ancestors of modern West Africans, bearing haplogroups [[Haplogroup E-M2#E1b1a1a1f|E1b1a1-L485]] and [[Haplogroup E-M2#E1b1a1a1g|E1b1a1-U175]] or their ancestral haplogroup E1b1a1-M4732.<ref name="Shriner" /> West Africans (e.g., [[Yoruba people|Yoruba]] and [[Esan people|Esan]] of Nigeria), bearing the Benin sickle cell haplotype, may have migrated through the [[Northeast Africa|northeastern region of Africa]] into the western region of [[Arabia]].<ref name="Shriner" /> West Africans (e.g., [[Mende people|Mende]] of Sierra Leone), bearing the Senegal sickle cell haplotype,<ref name="Yaseen">{{cite journal | vauthors = Yaseen NT, Al-Mamoori HS, Hassan MK |title=Sickle β-globin haplotypes among patients with sickle cell anemia in Basra, Iraq: A cross-sectional study | doi = 10.4103/ijh.ijh_20_19 |journal=Iraqi Journal of Hematology | date = January 2020 | volume = 9 | issue = 1 | pages = 23 |s2cid=216082225 }}</ref><ref name="Shriner" /> may have migrated into [[Mauritania]] (77% modern rate of occurrence) and Senegal (100%); they may also have migrated across the Sahara, into [[North Africa]], and from North Africa, into [[Southern Europe]], [[Turkey]], and a region near northern [[Iraq]] and southern Turkey.<ref name="Yaseen" /> Some may have migrated into and introduced the Senegal and Benin sickle cell haplotypes into [[Basra]], Iraq, where both occur equally.<ref name="Yaseen" /> West Africans, bearing the Benin sickle cell haplotype, may have migrated into the northern region of Iraq (69.5%), [[Jordan]] (80%), [[Lebanon]] (73%), [[Oman]] (52.1%), and [[Egypt]] (80.8%).<ref name="Yaseen" />
=====Schistosomes=====
According to Steverding (2020), while not definite: Near the [[African Great Lakes]], schistosomes (e.g., [[S. mansoni]], [[S. haematobium]]) underwent evolution.<ref name="Steverding">{{cite journal | vauthors = Steverding D | title = The spreading of parasites by human migratory activities | journal = Virulence | volume = 11 | issue = 1 | pages = 1177–1191 | date = December 2020 | pmid = 32862777 | pmc = 7549983 | doi = 10.1080/21505594.2020.1809963 }}</ref> Subsequently, there was an expansion alongside the Nile.<ref name="Steverding" /> From [[Egypt]], the presence of schistosomes may have expanded, via migratory [[Yoruba people]], into Western Africa.<ref name="Steverding" /> Thereafter, [[schistosomes]] may have expanded, via [[Bantu migration|migratory]] [[Bantu peoples]], into the rest of [[Sub-Saharan Africa]] (e.g., [[Southern Africa]], [[Central Africa]]).<ref name="Steverding" />
=====Thalassemia=====
Through pathways taken by [[Caravan (travellers)|caravan]]s, or via travel amid the [[Almoravid dynasty|Almovarid]] period, a population (e.g., [[Sub-Saharan]] [[West Africa#Demographics and languages|West Africans]]) may have introduced the –29 (A → G) [[Beta thalassemia|β-thalassemia]] mutation (found in notable amounts among [[African-Americans]]) into the [[North African]] region of [[Morocco]].<ref name="Agouti">{{cite journal | vauthors = Agouti I, Badens C, Abouyoub A, Levy N, Bennani M | title = Molecular basis of beta-thalassemia in Morocco: possible origins of the molecular heterogeneity | journal = Genetic Testing | volume = 12 | issue = 4 | pages = 563–568 | date = December 2008 | pmid = 18976160 | doi = 10.1089/gte.2008.0058 | s2cid = 46000591 }}</ref>
===Central Africa===
{{See also|African Pygmies#Genetics}}
====Archaic Human DNA====
{{Further|Interbreeding between archaic and modern humans#Archaic African hominins}}
Archaic traits found in human fossils of [[West Africa]] (e.g., [[Iwo Eleru skull|Iho Eleru fossils]], which dates to 13,000 BP) and [[Central Africa]] (e.g., [[Ishango]] fossils, which dates between 25,000 BP and 20,000 BP) may have developed as a result of admixture between archaic humans and modern humans or may be evidence of late-persisting [[Early modern human#Early Homo sapiens|early modern humans]].<ref name="Bergström"/> 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.<ref name="Bergström" />
====Ancient DNA====
In 4000 BP, there may have been a population that traversed from [[Africa]] (e.g., [[West Africa]] or West-[[Central Africa]]), through the [[Strait of Gibraltar]], into the [[Iberian peninsula]], where admixing between Africans and Iberians (e.g., of northern [[Portugal]], of southern [[Spain]]) occurred.<ref name="RSP"/>
=====Cameroon=====
[[West African hunter-gatherers]], in the region of western Central Africa (e.g., [[Shum Laka]], [[Cameroon]]), particularly between 8000 BP and 3000 BP, were found to be related to modern [[Central African foragers|Central African hunter-gatherers]] (e.g., [[Baka people (Cameroon and Gabon)|Baka]], [[Bakola]], [[Aka people|Biaka]], [[Bedzan people|Bedzan]]).<ref name="Lipson">{{cite journal | vauthors = Lipson M, Ribot I, Mallick S, Rohland N, Olalde I, Adamski N, Broomandkhoshbacht N, Lawson AM, López S, Oppenheimer J, Stewardson K, Asombang RN, Bocherens H, Bradman N, Culleton BJ, Cornelissen E, Crevecoeur I, de Maret P, Fomine FL, Lavachery P, Mindzie CM, Orban R, Sawchuk E, Semal P, Thomas MG, Van Neer W, Veeramah KR, Kennett DJ, Patterson N, Hellenthal G, Lalueza-Fox C, MacEachern S, Prendergast ME, Reich D | display-authors = 6 | title = Ancient West African foragers in the context of African population history | journal = Nature | volume = 577 | issue = 7792 | pages = 665–670 | date = January 2020 | pmid = 31969706 | pmc = 8386425 | doi = 10.1038/s41586-020-1929-1 | s2cid = 210862788 | bibcode = 2020Natur.577..665L }}</ref>
=====Democratic Republic of Congo=====
At Kindoki, in the [[Democratic Republic of Congo]], there were three individuals, dated to the [[protohistoric]] period (230 BP, 150 BP, 230 BP); one carried haplogroups [[Haplogroup E-M2#E1b1a1a1d|E1b1a1a1d1a2 (E-CTS99, E-CTS99)]] and [[Haplogroup L1 (mtDNA)#L1c|L1c3a1b]], another carried [[Haplogroup E-M96|haplogroup E (E-M96, E-PF1620)]], and the last carried haplogroups [[Haplogroup R1b#R1b (R-L278)|R1b1 (R-P25 1, R-M415)]] and [[Haplogroup L0 (mtDNA)#Distribution|L0a1b1a1]].<ref name="Wang">{{cite journal | vauthors = Wang K, Goldstein S, Bleasdale M, Clist B, Bostoen K, Bakwa-Lufu P, Buck LT, Crowther A, Dème A, McIntosh RJ, Mercader J, Ogola C, Power RC, Sawchuk E, Robertshaw P, Wilmsen EN, Petraglia M, Ndiema E, Manthi FK, Krause J, Roberts P, Boivin N, Schiffels S | display-authors = 6 | title = Ancient genomes reveal complex patterns of population movement, interaction, and replacement in sub-Saharan Africa | journal = Science Advances | volume = 6 | issue = 24 | pages = eaaz0183 | date = June 2020 | pmid = 32582847 | pmc = 7292641 | doi = 10.1126/sciadv.aaz0183 | bibcode = 2020SciA....6..183W }}</ref><ref name="Wang II">{{cite journal | vauthors = Wang K, Goldstein S, Bleasdale M, Clist B, Bostoen K, Bakwa-Lufu P, Buck LT, Crowther A, Dème A, McIntosh RJ, Mercader J, Ogola C, Power RC, Sawchuk E, Robertshaw P, Wilmsen EN, Petraglia M, Ndiema E, Manthi FK, Krause J, Roberts P, Boivin N, Schiffels S | display-authors = 6 | title = Ancient genomes reveal complex patterns of population movement, interaction, and replacement in sub-Saharan Africa | journal = Science Advances | volume = 6 | issue = 24 | pages = eaaz0183 | date = June 2020 | pmid = 32582847 | doi = 10.1126/sciadv.aaz0183 | pmc = 7292641 | bibcode = 2020SciA....6..183W }}</ref>
At Ngongo Mbata, in the [[Democratic Republic of Congo]], an individual, dated to the [[protohistoric]] period (220 BP), carried haplogroup [[Haplogroup L1 (mtDNA)#L1c|L1c3a]].<ref name="Wang" /><ref name="Wang II" />
At Matangai Turu Northwest, in the [[Democratic Republic of Congo]], an individual, dated to the [[Iron Age#Sub-Saharan Africa|Iron Age]] (750 BP), carried an undetermined haplogroup(s).<ref name="Wang" /><ref name="Wang II" />
====Y-Chromosomal DNA====
{{Further|Y-DNA haplogroups in populations of Sub-Saharan Africa}}
[[Haplogroup R1b#R1b1b (R-V88)|Haplogroup R-V88]] may have originated in western Central Africa (e.g., [[Equatorial Guinea]]), and, in the middle of the [[Holocene]], arrived in [[North Africa]] through population migration.<ref name="González-Santos">{{cite journal | vauthors = González M, Gomes V, López-Parra AM, Amorim A, Carracedo A, Sánchez-Diz P, Arroyo-Pardo E, Gusmão L | display-authors = 6 | title = The genetic landscape of Equatorial Guinea and the origin and migration routes of the Y chromosome haplogroup R-V88 | journal = European Journal of Human Genetics | volume = 21 | issue = 3 | pages = 324–331 | date = March 2013 | pmid = 22892526 | doi = 10.1038/ejhg.2012.167 | pmc = 3573200 }}</ref>
====Mitochondrial DNA====
{{Further|Haplogroup L1 (mtDNA)|Haplogroup L2 (mtDNA)}}
Mitochondrial [[Haplogroup L1 (mtDNA)|haplogroup L1c]] is strongly associated with pygmies, especially with [[Mbenga people|Bambenga]] groups.<ref name="Quintana08">{{cite journal | vauthors = Quintana-Murci L, Quach H, Harmant C, Luca F, Massonnet B, Patin E, Sica L, Mouguiama-Daouda P, Comas D, Tzur S, Balanovsky O, Kidd KK, Kidd JR, van der Veen L, Hombert JM, Gessain A, Verdu P, Froment A, Bahuchet S, Heyer E, Dausset J, Salas A, Behar DM | display-authors = 6 | title = Maternal traces of deep common ancestry and asymmetric gene flow between Pygmy hunter-gatherers and Bantu-speaking farmers | journal = Proceedings of the National Academy of Sciences of the United States of America | volume = 105 | issue = 5 | pages = 1596–1601 | date = February 2008 | pmid = 18216239 | pmc = 2234190 | doi = 10.1073/pnas.0711467105 | doi-access = free | bibcode = 2008PNAS..105.1596Q }}</ref> L1c prevalence was variously reported as: 100% in [[Gyele language|Ba-Kola]], 97% in [[Aka people|Aka (Ba-Benzélé)]], and 77% in [[Aka people|Biaka]],<ref name="Tishkoff">Sarah A. Tishkoff et al. 2007, [https://web.archive.org/web/20080517050807/http://mbe.oxfordjournals.org/cgi/content/full/24/10/2180 History of Click-Speaking Populations of Africa Inferred from mtDNA and Y Chromosome Genetic Variation.] Molecular Biology and Evolution 2007 24(10):2180-2195</ref> 100% of the [[Bedzan people|Bedzan (Tikar)]], 97% and 100% in the [[Baka (Cameroon and Gabon)|Baka people]] of [[Gabon]] and [[Cameroon]], respectively,<ref>Lluis Quintana-Murci et al. MtDNA diversity in Central Africa: from hunter-gathering to agriculturalism. CNRS-Institut Pasteur, Paris</ref> 97% in [[Bakoya]] (97%), and 82% in [[Bongo people (Gabon)|Ba-Bongo]].<ref name="Quintana08" /> Mitochondrial haplogroups [[Haplogroup L2 (mtDNA)|L2a]] and [[Haplogroup L0 (mtDNA)|L0a]] are prevalent among the [[Bambuti]].<ref name="Quintana08"/><ref>{{cite journal | vauthors = Silva M, Alshamali F, Silva P, Carrilho C, Mandlate F, Jesus Trovoada M, Černý V, Pereira L, Soares P | display-authors = 6 | title = 60,000 years of interactions between Central and Eastern Africa documented by major African mitochondrial haplogroup L2 | journal = Scientific Reports | volume = 5 | pages = 12526 | date = July 2015 | pmid = 26211407 | pmc = 4515592 | doi = 10.1038/srep12526 | bibcode = 2015NatSR...512526S }}</ref>
====Autosomal DNA====
Genetically, [[African pygmies]] have some key difference between them and [[Bantu peoples]].<ref>{{cite journal | vauthors = Jarvis JP, Scheinfeldt LB, Soi S, Lambert C, Omberg L, Ferwerda B, Froment A, Bodo JM, Beggs W, Hoffman G, Mezey J, Tishkoff SA | display-authors = 6 | title = Patterns of ancestry, signatures of natural selection, and genetic association with stature in Western African pygmies | journal = PLOS Genetics | volume = 8 | issue = 4 | pages = e1002641 | year = 2012 | pmid = 22570615 | pmc = 3343053 | doi = 10.1371/journal.pgen.1002641 }}</ref><ref>{{cite journal | vauthors = López Herráez D, Bauchet M, Tang K, Theunert C, Pugach I, Li J, Nandineni MR, Gross A, Scholz M, Stoneking M | display-authors = 6 | title = Genetic variation and recent positive selection in worldwide human populations: evidence from nearly 1 million SNPs | journal = PLOS ONE | volume = 4 | issue = 11 | pages = e7888 | date = November 2009 | pmid = 19924308 | pmc = 2775638 | doi = 10.1371/journal.pone.0007888 | doi-access = free | bibcode = 2009PLoSO...4.7888L }}</ref>
====Medical DNA====
Evidence suggests that, when compared to other [[Sub-Saharan]] African populations, [[African pygmy]] populations display unusually low levels of expression of the genes encoding for [[human growth hormone]] and [[Growth hormone receptor|its receptor]] associated with low [[Blood serum|serum]] levels of [[insulin-like growth factor-1]] and short stature.<ref name='Bozzola, 2009'>{{cite journal | vauthors = Bozzola M, Travaglino P, Marziliano N, Meazza C, Pagani S, Grasso M, Tauber M, Diegoli M, Pilotto A, Disabella E, Tarantino P, Brega A, Arbustini E | display-authors = 6 | title = The shortness of Pygmies is associated with severe under-expression of the growth hormone receptor | journal = Molecular Genetics and Metabolism | volume = 98 | issue = 3 | pages = 310–313 | date = November 2009 | pmid = 19541519 | doi = 10.1016/j.ymgme.2009.05.009 }}</ref>
===Eastern Africa===
====Archaic Human DNA====
{{Further|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.<ref name="Bergström"/>
====Ancient DNA====
=====Ethiopia=====
At [[Mota, Ethiopia|Mota]], in [[Ethiopia]], an individual, estimated to date to the 5th millennium BP, carried haplogroups [[Haplogroup E-P2|E1b1]] and [[Haplogroup L3 (mtDNA)#Subclade distribution|L3x2a]].<ref name="Llorente">{{cite journal | vauthors = Llorente MG, Jones ER, Eriksson A, Siska V, Arthur KW, Arthur JW, Curtis MC, Stock JT, Coltorti M, Pieruccini P, Stretton S, Brock F, Higham T, Park Y, Hofreiter M, Bradley DG, Bhak J, Pinhasi R, Manica A | display-authors = 6 | title = Ancient Ethiopian genome reveals extensive Eurasian admixture throughout the African continent | journal = Science | volume = 350 | issue = 6262 | pages = 820–822 | date = November 2015 | pmid = 26449472 | doi = 10.1126/science.aad2879 | bibcode = 2015Sci...350..820L | hdl = 2318/1661894 | s2cid = 25743789 }}</ref><ref name="Llorente II">{{cite journal | vauthors = Llorente MG, Jones ER, Eriksson A, Siska V, Arthur KW, Arthur JW, Curtis MC, Stock JT, Coltorti M, Pieruccini P, Stretton S, Brock F, Higham T, Park Y, Hofreiter M, Bradley DG, Bhak J, Pinhasi R, Manica A | display-authors = 6 |title=Supplementary Materials for Ancient Ethiopian genome reveals extensive Eurasian admixture in Eastern Africa |url=https://www.science.org/doi/10.1126/science.aad2879 |journal=Science | date = 13 November 2015 | volume = 350 | issue = 6262 | pages = 820–822 | doi = 10.1126/science.aad2879 | pmid = 26449472 | bibcode = 2015Sci...350..820L | hdl = 2318/1661894 | s2cid = 25743789 }}</ref> 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.<ref name="Hellenthal">{{cite journal | vauthors = Hellenthal G, Bird N, Morris S | title = Structure and ancestry patterns of Ethiopians in genome-wide autosomal DNA | journal = Human Molecular Genetics | volume = 30 | issue = R1 | pages = R42–R48 | date = April 2021 | pmid = 33547782 | pmc = 8242491 | doi = 10.1093/hmg/ddab019 }}</ref><ref>{{Cite journal |last1=Pagani |first1=Luca |last2=Kivisild |first2=Toomas |last3=Tarekegn |first3=Ayele |last4=Ekong |first4=Rosemary |last5=Plaster |first5=Chris |last6=Gallego Romero |first6=Irene |last7=Ayub |first7=Qasim |last8=Mehdi |first8=S. Qasim |last9=Thomas |first9=Mark G. |last10=Luiselli |first10=Donata |last11=Bekele |first11=Endashaw |date=2012-07-13 |title=Ethiopian genetic diversity reveals linguistic stratification and complex influences on the Ethiopian gene pool |journal=American Journal of Human Genetics |volume=91 |issue=1 |pages=83–96 |doi=10.1016/j.ajhg.2012.05.015 |issn=1537-6605 |pmc=3397267 |pmid=22726845}}</ref>
=====Kenya=====
At Jawuoyo Rockshelter, in [[Kisumu County]], [[Kenya]], a forager of the Later [[Stone Age]] carried haplogroups [[Haplogroup E-V68#E-V22|E1b1b1a1b2/E-V22]] and [[Haplogroup L4 (mtDNA)|L4b2a2c]].<ref name="Prendergast">{{cite journal | vauthors = Prendergast ME, Lipson M, Sawchuk EA, Olalde I, Ogola CA, Rohland N, Sirak KA, Adamski N, Bernardos R, Broomandkhoshbacht N, Callan K, Culleton BJ, Eccles L, Harper TK, Lawson AM, Mah M, Oppenheimer J, Stewardson K, Zalzala F, Ambrose SH, Ayodo G, Gates HL, Gidna AO, Katongo M, Kwekason A, Mabulla AZ, Mudenda GS, Ndiema EK, Nelson C, Robertshaw P, Kennett DJ, Manthi FK, Reich D | display-authors = 6 | title = Ancient DNA reveals a multistep spread of the first herders into sub-Saharan Africa | journal = Science | volume = 365 | issue = 6448 | date = July 2019 | pmid = 31147405 | doi = 10.1126/science.aaw6275 | pmc = 6827346 | bibcode = 2019Sci...365.6275P }}</ref><ref name="Prendergast II">{{cite journal | vauthors = Prendergast ME, Lipson M, Sawchuk EA, Olalde I, Ogola CA, Rohland N, Sirak KA, Adamski N, Bernardos R, Broomandkhoshbacht N, Callan K, Culleton BJ, Eccles L, Harper TK, Lawson AM, Mah M, Oppenheimer J, Stewardson K, Zalzala F, Ambrose SH, Ayodo G, Gates HL, Gidna AO, Katongo M, Kwekason A, Mabulla AZ, Mudenda GS, Ndiema EK, Nelson C, Robertshaw P, Kennett DJ, Manthi FK, Reich D | display-authors = 6 |title=Supplementary Materials for Ancient DNA reveals a multistep spread of the first herders into sub-Saharan Africa |journal=Science | date = 5 July 2019 | volume = 365 | issue = 6448 | pages = eaaw6275 | doi = 10.1126/science.aaw6275 | pmid = 31147405 | pmc = 6827346 | bibcode = 2019Sci...365.6275P }}</ref>
At Ol Kalou, in [[Nyandarua County]], [[Kenya]], a pastoralist of the [[Pastoral Neolithic]] carried haplogroups [[Haplogroup E-Z827#E-M293|E1b1b1b2b2a1/E-M293]] and [[Haplogroup L3 (mtDNA)#Subclade distribution|L3d1d]].<ref name="Prendergast" /><ref name="Prendergast II" />
At Kokurmatakore, in [[Marsabit County]], [[Kenya]], a pastoralist of the Pastoral [[Iron Age#Sub-Saharan Africa|Iron Age]] carried haplogroups [[Haplogroup E-M35|E1b1b1/E-M35]] and [[Haplogroup L3 (mtDNA)#Subclade distribution|L3a2a]].<ref name="Prendergast" /><ref name="Prendergast II" />
At White Rock Point, in [[Homa Bay County]], [[Kenya]], there were two foragers of the Later [[Stone Age]]; one carried haplogroups [[Haplogroup BT|BT (xCT)]], likely [[Haplogroup B-M60|B]], and [[Haplogroup L2 (mtDNA)#Haplogroup L2a|L2a4]], and another probably carried [[Haplogroup L0 (mtDNA)#Distribution|haplogroup L0a2]].<ref name="Prendergast" /><ref name="Prendergast II" />
At Nyarindi Rockshelter, in [[Kenya]], there were two individuals, dated to the Later [[Stone Age]] (3500 BP); one carried [[Haplogroup L4 (mtDNA)|haplogroup L4b2a]] and another carried [[Haplogroup E-M96|haplogroup E (E-M96, E-P162)]].<ref name="Wang" /><ref name="Wang II" />
At Lukenya Hill, in [[Kenya]], there were two individuals, dated to the [[Pastoral Neolithic]] (3500 BP); one carried haplogroups [[Haplogroup E-Z827#E-V1515|E1b1b1b2b (E-M293, E-CTS10880)]] and [[Haplogroup L4 (mtDNA)|L4b2a2b]], and another carried [[Haplogroup L0 (mtDNA)#Distribution|haplogroup L0f1]].<ref name="Wang" /><ref name="Wang II" />
At Hyrax Hill, in [[Kenya]], an individual, dated to the [[Pastoral Neolithic]] (2300 BP), carried haplogroups [[Haplogroup E-Z827#E-V1515|E1b1b1b2b (E-M293, E-M293)]] and [[Haplogroup L5 (mtDNA)|L5a1b]].<ref name="Wang" /><ref name="Wang II" />
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 [[Haplogroup E-Z827#E-V1515|E1b1b1b2b (E-M293, E-M293)]] and [[Haplogroup L3 (mtDNA)#Subclade distribution|L3h1a2a1]].<ref name="Wang" /><ref name="Wang II" />
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#Sub-Saharan Africa|Iron Age]] (300 BP, 900 BP); one carried haplogroups [[Haplogroup CT|CT (CT-M168, CT-M5695)]] and [[Haplogroup L3 (mtDNA)#Subclade distribution|L3i1]], another carried [[Haplogroup L2 (mtDNA)#Haplogroup L2a1|haplogroup L2a1f]], and the last carried [[Haplogroup L2 (mtDNA)#Haplogroup L2a|haplogroup L2a5]].<ref name="Wang" /><ref name="Wang II" />
At [[Panga ya Saidi]], in [[Kenya]], an individual, estimated to date between 496 BP and 322 BP, carried haplogroups [[Haplogroup E-M215 (Y-DNA)#E-Z830 (E1b1b1b2)|E1b1b1b2]] and [[Haplogroup L4 (mtDNA)|L4b2a2]].<ref name="Skoglund">{{cite journal | vauthors = Skoglund P, Thompson JC, Prendergast ME, Mittnik A, Sirak K, Hajdinjak M, Salie T, Rohland N, Mallick S, Peltzer A, Heinze A, Olalde I, Ferry M, Harney E, Michel M, Stewardson K, Cerezo-Román JI, Chiumia C, Crowther A, Gomani-Chindebvu E, Gidna AO, Grillo KM, Helenius IT, Hellenthal G, Helm R, Horton M, López S, Mabulla AZ, Parkington J, Shipton C, Thomas MG, Tibesasa R, Welling M, Hayes VM, Kennett DJ, Ramesar R, Meyer M, Pääbo S, Patterson N, Morris AG, Boivin N, Pinhasi R, Krause J, Reich D | display-authors = 6 | title = Reconstructing Prehistoric African Population Structure | journal = Cell | volume = 171 | issue = 1 | pages = 59–71.e21 | date = September 2017 | pmid = 28938123 | pmc = 5679310 | doi = 10.1016/j.cell.2017.08.049 }}</ref>
======Laikipia County======
At Kisima Farm/Porcupine Cave, in [[Laikipia County]], [[Kenya]], there were two pastoralists of the [[Pastoral Neolithic]]; one carried haplogroups [[Haplogroup E-Z827#E-M293|E1b1b1b2b2a1/E-M293]] and [[Haplogroup M (mtDNA)#Haplogroup M1|M1a1]], and another carried haplogroup [[Haplogroup M (mtDNA)#Haplogroup M1|M1a1f]].<ref name="Prendergast" /><ref name="Prendergast II" />
At Kisima Farm/C4, in [[Laikipia County]], [[Kenya]], a pastoralist of the Pastoral [[Iron Age#Sub-Saharan Africa|Iron Age]], carried haplogroups [[Haplogroup E-M75|E2 (xE2b)/E-M75]] and [[Haplogroup L3 (mtDNA)#Subclade distribution|L3h1a1]].<ref name="Prendergast" /><ref name="Prendergast II" />
At Laikipia District Burial, in [[Laikipia County]], [[Kenya]], a pastoralist of the Pastoral [[Iron Age#Sub-Saharan Africa|Iron Age]] carried haplogroup [[Haplogroup L0 (mtDNA)#Distribution|L0a1c1]].<ref name="Prendergast" /><ref name="Prendergast II" />
======Nakuru County======
At Prettejohn's Gully, in [[Nakuru County, Kenya]], there were two pastoralists of the early pastoral period; one carried haplogroups [[Haplogroup E-M75|E2 (xE2b)/E-M75]] and [[Haplogroup K (mtDNA)|K1a]], and another carried haplogroup [[Haplogroup L3 (mtDNA)#Subclade distribution|L3f1b]].<ref name="Prendergast" /><ref name="Prendergast II" />
At Cole's Burial, in [[Nakuru County, Kenya]], a pastoralist of the [[Pastoral Neolithic]] carried haplogroups [[Haplogroup E-V68#Subclades of M78|E1b1b1a1a1b1/E-CTS3282]] and [[Haplogroup L3 (mtDNA)#Subclade distribution|L3i2]].<ref name="Prendergast" /><ref name="Prendergast II" />
At Rigo Cave, in [[Nakuru County, Kenya]], there were three pastoralists of the [[Pastoral Neolithic]]/[[Elmenteitan]], one carried haplogroups [[Haplogroup E-Z827#E-M293|E1b1b1b2b2a1/E-M293]] and [[Haplogroup L3 (mtDNA)#Subclade distribution|L3f]], another carried haplogroups [[Haplogroup E-Z827#E-V1515|E1b1b1b2b2/E-V1486]], likely [[Haplogroup E-M215 (Y-DNA)#E-M293|E-M293]], and probably [[Haplogroup M (mtDNA)#Haplogroup M1|M1a1b]], and the last carried haplogroups [[Haplogroup E-M215 (Y-DNA)#E-M293|E1b1b1b2b2a1/E-M293]] and [[Haplogroup L4 (mtDNA)|L4b2a2c]].<ref name="Prendergast" /><ref name="Prendergast II" />
At Naishi Rockshelter, in [[Nakuru County, Kenya]], there two pastoralists of the [[Pastoral Neolithic]]; one carried haplogroups [[Haplogroup E-Z827#E-V1515|E1b1b1b2b/E-V1515]], likely [[Haplogroup E-M215 (Y-DNA)#E-M293|E-M293]], and [[Haplogroup L3 (mtDNA)#Subclade distribution|L3x1a]], and another carried haplogroups [[Haplogroup A (Y-DNA)#Structure & subclade distribution|A1b (xA1b1b2a)/A-P108]] and [[Haplogroup L0 (mtDNA)#Distribution|L0a2d]].<ref name="Prendergast" /><ref name="Prendergast II" />
At Keringet Cave, in [[Nakuru County, Kenya]], a pastoralist of the [[Pastoral Neolithic]] carried [[Haplogroup A (Y-DNA)#Structure & subclade distribution|haplogroups A1b1b2/A-L427]] and [[Haplogroup L4 (mtDNA)|L4b2a1]], and another pastoralist of the [[Pastoral Neolithic]]/[[Elmenteitan]] carried [[Haplogroup K (mtDNA)|haplogroup K1a]].<ref name="Prendergast" /><ref name="Prendergast II" />
At Naivasha Burial Site, in [[Nakuru County, Kenya]], there were five pastoralists of the [[Pastoral Neolithic]]; one carried [[Haplogroup L4 (mtDNA)|haplogroup L4b2a2b]], another carried haplogroups xBT, likely [[Haplogroup A (Y-DNA)|A]], and [[Haplogroup M (mtDNA)#Haplogroup M1|M1a1b]], another carried haplogroups [[Haplogroup E-Z827#E-M293|E1b1b1b2b2a1/E-M293]] and [[Haplogroup L3 (mtDNA)#Subclade distribution|L3h1a1]], another carried haplogroups [[Haplogroup A (Y-DNA)#A1b1b2b (A-M13)|A1b1b2b/A-M13]] and [[Haplogroup L4 (mtDNA)|L4a1]], and the last carried haplogroups [[Haplogroup E-Z827#E-M293|E1b1b1b2b2a1/E-M293]] and [[Haplogroup L3 (mtDNA)#Subclade distribution|L3x1a]].<ref name="Prendergast" /><ref name="Prendergast II" />
At Njoro River Cave II, in [[Nakuru County, Kenya]], a pastoralist of the [[Pastoral Neolithic]] carried haplogroup [[Haplogroup L3 (mtDNA)#Subclade distribution|L3h1a2a1]].<ref name="Prendergast" /><ref name="Prendergast II" />
At Egerton Cave, in [[Nakuru County, Kenya]], a pastoralist of the [[Pastoral Neolithic]]/[[Elmenteitan]] carried haplogroup [[Haplogroup L0 (mtDNA)#Distribution|L0a1d]].<ref name="Prendergast" /><ref name="Prendergast II" />
At Ilkek Mounds, in [[Nakuru County, Kenya]], a pastoralist of the Pastoral [[Iron Age#Sub-Saharan Africa|Iron Age]] carried haplogroups [[Haplogroup E-M75|E2 (xE2b)/E-M75]] and [[Haplogroup L0 (mtDNA)#Distribution|L0f2a]].<ref name="Prendergast" /><ref name="Prendergast II" />
At Deloraine Farm, in [[Nakuru County, Kenya]], an iron metallurgist of the [[Iron Age#Sub-Saharan Africa|Iron Age]] carried haplogroups [[Haplogroup E-M2#E1b1a1a1a|E1b1a1a1a1a/E-M58]] and [[Haplogroup L5 (mtDNA)|L5b1]].<ref name="Prendergast" /><ref name="Prendergast II" />
======Narok County======
At Kasiole 2, in [[Narok County]], [[Kenya]], a pastoralist of the Pastoral [[Iron Age#Sub-Saharan Africa|Iron Age]] carried haplogroups [[Haplogroup E-Z827#E-V1515|E1b1b1b2b/E-V1515]], likely [[Haplogroup E-Z827#E-M293|E-M293]], and [[Haplogroup L3 (mtDNA)#Subclade distribution|L3h1a2a1]].<ref name="Prendergast" /><ref name="Prendergast II" />
At Emurua Ole Polos, in [[Narok County]], [[Kenya]], a pastoralist of the Pastoral [[Iron Age#Sub-Saharan Africa|Iron Age]] carried haplogroups [[Haplogroup E-Z827#E-M293|E1b1b1b2b2a1/E-M293]] and [[Haplogroup L3 (mtDNA)#Subclade distribution|L3h1a2a1]].<ref name="Prendergast" /><ref name="Prendergast II" />
=====Tanzania=====
At [[Luxmanda]], [[Tanzania]], an individual, estimated to date between 3141 BP and 2890 BP, carried [[Haplogroup L2 (mtDNA)#Haplogroup L2a1|haplogroup L2a1]].<ref name="Skoglund" />
At [[Kuumbi Cave, Zanzibar|Kuumbi Cave]], in [[Zanzibar]], [[Tanzania]], an individual, estimated to date between 1370 BP and 1303 BP, carried haplogroup [[Haplogroup L4 (mtDNA)|L4b2a2c]].<ref name="Skoglund" />
======Karatu District======
At Gishimangeda Cave, in [[Karatu District]], [[Tanzania]], there were eleven pastoralists of the [[Pastoral Neolithic]]; one carried haplogroups [[Haplogroup E-V68#Family tree|E1b1b1a1b2/E-V22]] and [[Haplogroup HV (mtDNA)#Tree|HV1b1]], another carried [[Haplogroup L0 (mtDNA)#Distribution|haplogroup L0a]], another carried haplogroup [[Haplogroup L3 (mtDNA)#Subclade distribution|L3x1]], another carried haplogroup [[Haplogroup L4 (mtDNA)|L4b2a2b]], another carried haplogroups [[Haplogroup E-Z827#E-M293|E1b1b1b2b2a1/E-M293]] and [[Haplogroup L3 (mtDNA)#Subclade distribution|L3i2]], another carried haplogroup [[Haplogroup L3 (mtDNA)#Subclade distribution|L3h1a2a1]], another carried haplogroups [[Haplogroup E-Z827#E-V1515|E1b1b1b2b2/E-V1486]], likely [[Haplogroup E-Z827#E-M293|E-M293]] and [[Haplogroup L0 (mtDNA)#Distribution|L0f2a1]], and another carried haplogroups [[Haplogroup E-Z827#E-V1515|E1b1b1b2b2/E-V1486]], likely [[Haplogroup E-Z827#E-M293|E-M293]], and [[Haplogroup T (mtDNA)|T2+150]]; while most of the haplogroups among three pastoralists went undetermined, one was determined to carry [[haplogroup BT]], likely [[Haplogroup B-M60|B]].<ref name="Prendergast" /><ref name="Prendergast II" />
======Pemba Island======
At Makangale Cave, on [[Pemba Island]], [[Tanzania]], an individual, estimated to date between 1421 BP and 1307 BP, carried [[Haplogroup L0 (mtDNA)#Distribution|haplogroup L0a]].<ref name="Skoglund" />
At Makangale Cave, on [[Pemba Island]], [[Tanzania]], an individual, estimated to date between 639 BP and 544 BP, carried [[Haplogroup L2 (mtDNA)#Haplogroup L2a1a|haplogroup L2a1a2]].<ref name="Skoglund" />
=====Uganda=====
At [[Munsa]], in [[Uganda]], an individual, dated to the Later [[Iron Age#Sub-Saharan Africa|Iron Age]] (500 BP), carried haplogroup [[Haplogroup L3 (mtDNA)#Subclade distribution|L3b1a1]].<ref name="Wang" /><ref name="Wang II" />
====Y-Chromosomal DNA====
{{Further|Y-DNA haplogroups in populations of Sub-Saharan Africa}}
As of 19,000 years ago, Africans, bearing [[Haplogroup E-V38|haplogroup E1b1a-V38]], likely traversed across the [[Sahara]], from [[East Africa|east]] to [[West Africa|west]].<ref name="Shriner"/>
Before the [[Trans-Saharan slave trade#Trans-Saharan slave trade in the Middle Ages|slave]] [[Indian Ocean slave trade#Muslim Period|trade]] period, East Africans, who carried [[Haplogroup E-M2|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]]).<ref name="Fernandes">{{cite web | vauthors = Neves da Nova Fernandes VC |title=High-resolution characterization of genetic markers in the Arabian Peninsula and Near East |url=https://etheses.whiterose.ac.uk/6301/1/Vfernandes_PhD_thesis.pdf |website=White Rose eTheses Online |publisher=University of Leeds}}</ref>
====Mitochondrial DNA====
{{Further|Macro-haplogroup L (mtDNA)|Haplogroup M (mtDNA)|Haplogroup N (mtDNA)}}
====Autosomal DNA====
Across all areas of [[Madagascar]], the average ancestry for the [[Malagasy people]] was found to be 4% [[West Eurasian]], 37% [[Austronesian peoples|Austronesian]], and 59% [[Bantu peoples|Bantu]].<ref name="Heiske">{{cite journal | vauthors = Heiske M, Alva O, Pereda-Loth V, Van Schalkwyk M, Radimilahy C, Letellier T, Rakotarisoa JA, Pierron D | display-authors = 6 | title = Genetic evidence and historical theories of the Asian and African origins of the present Malagasy population | journal = Human Molecular Genetics | volume = 30 | issue = R1 | pages = R72–R78 | date = April 2021 | pmid = 33481023 | doi = 10.1093/hmg/ddab018 | doi-access = free }}</ref>
====Medical DNA====
===Southern Africa===
{{See also|San people#Genetics|Khoekhoe}}
====Archaic Human DNA====
{{Further|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.<ref name="Bergström"/>
====Ancient DNA====
Three [[Later Stone Age]] [[hunter-gatherers]] carried ancient DNA similar to [[Khoisan]]-speaking hunter-gatherers.<ref name="Choudhury">{{cite journal | vauthors = Choudhury A, Sengupta D, Ramsay M, Schlebusch C | title = Bantu-speaker migration and admixture in southern Africa | journal = Human Molecular Genetics | volume = 30 | issue = R1 | pages = R56–R63 | date = April 2021 | pmid = 33367711 | pmc = 8117461 | doi = 10.1093/hmg/ddaa274 }}</ref> Prior to the [[Bantu migration]] into the region, as evidenced by ancient DNA from [[Botswana]], [[East Africa]]n [[herders]] migrated into Southern Africa.<ref name="Choudhury" /> Out of four [[Iron Age]] Bantu [[agriculturalists]] of [[West African]] origin, two earlier agriculturalists carried ancient DNA similar to [[Tsonga people|Tsonga]] and [[Venda people|Venda]] peoples and the two later agriculturalists carried ancient DNA similar to [[Nguni people]]; this indicates that there were various movements of peoples in the overall Bantu migration, which resulted in increased interaction and admixing between [[Bantu languages|Bantu-speaking]] peoples and Khoisan-speaking peoples.<ref name="Choudhury" />
=====Botswana=====
At Nqoma, in [[Botswana]], an individual, dated to the Early [[Iron Age#Sub-Saharan Africa|Iron Age]] (900 BP), carried [[Haplogroup L2 (mtDNA)#Haplogroup L2a1|haplogroup L2a1f]].<ref name="Wang" /><ref name="Wang II" />
At Taukome, in [[Botswana]], an individual, dated to the Early [[Iron Age#Sub-Saharan Africa|Iron Age]] (1100 BP), carried haplogroups [[Haplogroup E-M2#E1b1a1|E1b1a1 (E-M2, E-Z1123)]] and [[Haplogroup L0 (mtDNA)#Distribution|L0d3b1]].<ref name="Wang" /><ref name="Wang II" />
At Xaro, in [[Botswana]], there were two individuals, dated to the Early [[Iron Age#Sub-Saharan Africa|Iron Age]] (1400 BP); one carried haplogroups [[Haplogroup E-M2#E1b1a1a1c|E1b1a1a1c1a]] and [[Haplogroup L3 (mtDNA)#Subclade distribution|L3e1a2]], and another carried haplogroups [[Haplogroup E-Z827#E-V1515|E1b1b1b2b (E-M293, E-CTS10880)]] and [[Haplogroup L0 (mtDNA)#Distribution|L0k1a2]].<ref name="Wang" /><ref name="Wang II" />
=====Malawi=====
======Fingira======
At Fingira, in [[Malawi]], an individual, estimated to date between 6175 BP and 5913 BP, carried haplogroups [[Haplogroup BT|BT]] and [[Haplogroup L0 (mtDNA)#Distribution|L0d1b2b]].<ref name="Skoglund" />
At Fingira, in [[Malawi]], an individual, estimated to date between 6177 BP and 5923 BP, carried haplogroups [[Haplogroup BT|BT]] and [[Haplogroup L0 (mtDNA)#Distribution|L0d1c]].<ref name="Skoglund" />
At Fingira, in [[Malawi]], an individual, estimated to date between 2676 BP and 2330 BP, carried [[Haplogroup L0 (mtDNA)#Distribution|haplogroup L0f]].<ref name="Skoglund" />
======Chencherere======
At Chencherere, in [[Malawi]], an individual, estimated to date between 5400 BP and 4800 BP, carried [[Haplogroup L0 (mtDNA)#Distribution|haplogroup L0k2]].<ref name="Skoglund" />
At Chencherere, in [[Malawi]], an individual, estimated to date between 5293 BP and 4979 BP, carried [[Haplogroup L0 (mtDNA)#Distribution|haplogroup L0k1]].<ref name="Skoglund" />
======Hora======
At Hora, in [[Malawi]], an individual, estimated to date between 10,000 BP and 5000 BP, carried haplogroups [[Haplogroup BT|BT]] and [[Haplogroup L0 (mtDNA)#Distribution|L0k2]].<ref name="Skoglund" />
At Hora, in [[Malawi]], an individual, estimated to date between 8173 BP and 7957 BP, carried [[Haplogroup L0 (mtDNA)#Distribution|haplogroup L0a2]].<ref name="Skoglund" />
=====South Africa=====
At [[Doonside, KwaZulu-Natal|Doonside]], in [[South Africa]], an individual, estimated to date between 2296 BP and 1910 BP, carried [[Haplogroup L0 (mtDNA)#Distribution|haplogroup L0d2]].<ref name="Schlebusch" /><ref name="Schlebusch II" />
At [[Champagne Castle]], in [[South Africa]], an individual, estimated to date between 448 BP and 282 BP, carried [[Haplogroup L0 (mtDNA)#Distribution|haplogroup L0d2a1a]].<ref name="Schlebusch" /><ref name="Schlebusch II" />
At [[Elands Bay Cave|Eland Cave]], in [[South Africa]], an individual, estimated to date between 533 BP and 453 BP, carried [[Haplogroup L3 (mtDNA)#Subclade distribution|haplogroup L3e3b1]].<ref name="Schlebusch" /><ref name="Schlebusch II" />
At Mfongosi, in [[South Africa]], an individual, estimated to date between 448 BP and 308 BP, carried [[Haplogroup L3 (mtDNA)#Subclade distribution|haplogroup L3e1b2]].<ref name="Schlebusch" /><ref name="Schlebusch II" />
At [[Newcastle, KwaZulu-Natal|Newcastle]], in [[South Africa]], an individual, estimated to date between 508 BP and 327 BP, carried [[Haplogroup L3 (mtDNA)#Subclade distribution|haplogroup L3e2b1a2]].<ref name="Schlebusch" /><ref name="Schlebusch II" />
At [[St Helena Bay|St. Helena]], in [[South Africa]], an individual, estimated to date between 2241 BP and 1965 BP, carried haplogroups [[Haplogroup A (Y-DNA)#A1b1b2a (A-M51)|A1b1b2a]] and [[Haplogroup L0 (mtDNA)#Distribution|L0d2c1]].<ref name="Skoglund" />
At Faraoskop Rock Shelter, in [[South Africa]], an individual, estimated to date between 2017 BP and 1748 BP, carried haplogroups [[Haplogroup A (Y-DNA)#A1b1b2a (A-M51)|A1b1b2a]] and [[Haplogroup L0 (mtDNA)#Distribution|L0d1b2b1b]].<ref name="Skoglund" />
At Kasteelberg, in [[South Africa]], an individual, estimated to date between 1282 BP and 1069 BP, carried haplogroup [[Haplogroup L0 (mtDNA)#Distribution|L0d1a1a]].<ref name="Skoglund" />
At Vaalkrans Shelter, in [[South Africa]], an individual, estimated to date to 200 BP, is predominantly related to [[Khoisan]] speakers, partly related (15% - 32%) to [[East Africa#Demographics|East Africans]], and carried haplogroups [[Haplogroup L0 (mtDNA)#Distribution|L0d3b1]].<ref name="Coutinho">{{cite journal | vauthors = Coutinho A, Malmström H, Edlund H, Henshilwood CS, van Niekerk KL, Lombard M, Schlebusch CM, Jakobsson M | display-authors = 6 | title = Later Stone Age human hair from Vaalkrans Shelter, Cape Floristic Region of South Africa, reveals genetic affinity to Khoe groups | journal = American Journal of Physical Anthropology | volume = 174 | issue = 4 | pages = 701–713 | date = April 2021 | pmid = 33539553 | doi = 10.1002/ajpa.24236 | publisher = Am J Phys Anthropol | s2cid = 213563734 }}</ref>
======Ballito Bay======
At [[Ballito]] Bay, [[South Africa]], an individual, estimated to date between 1986 BP and 1831 BP, carried haplogroups [[Haplogroup A (Y-DNA)#Structure & subclade distribution|A1b1b2]] and [[Haplogroup L0 (mtDNA)#Distribution|L0d2c1]].<ref name="Schlebusch">{{cite journal | vauthors = Schlebusch CM, Malmström H, Günther T, Sjödin P, Coutinho A, Edlund H, Munters AR, Vicente M, Steyn M, Soodyall H, Lombard M, Jakobsson M | display-authors = 6 | title = Southern African ancient genomes estimate modern human divergence to 350,000 to 260,000 years ago | journal = Science | volume = 358 | issue = 6363 | pages = 652–655 | date = November 2017 | pmid = 28971970 | doi = 10.1126/science.aao6266 | s2cid = 206663925 | doi-access = free | bibcode = 2017Sci...358..652S }}</ref><ref name="Schlebusch II">{{cite journal | vauthors = Schlebusch CM, Malmström H, Günther T, Sjödin P, Coutinho A, Edlund H, Munters AR, Vicente M, Steyn M, Soodyall H, Lombard M, Jakobsson M | display-authors = 6 |title=Supplementary Materials for Southern African ancient genomes estimate modern human divergence to 350,000to 260,000years ago |url=https://www.science.org/doi/10.1126/science.aao6266 |journal=Science | date = 3 November 2017 | volume = 358 | issue = 6363 | pages = 652–655 | doi = 10.1126/science.aao6266 | pmid = 28971970 | bibcode = 2017Sci...358..652S | s2cid = 206663925 }}</ref>
At [[Ballito]] Bay, [[South Africa]], an individual, estimated to date between 2149 BP and 1932 BP, carried haplogroups [[Haplogroup A (Y-DNA)#Structure & subclade distribution|A1b1b2]] and [[Haplogroup L0 (mtDNA)#Distribution|L0d2a1]].<ref name="Schlebusch" /><ref name="Schlebusch II" />
====Y-Chromosomal DNA====
{{Further|Y-DNA haplogroups in populations of Sub-Saharan Africa}}
[[File:Distribution of Y-Chromosome Haplogroup A in Africa.png|thumb|Distribution of Y-Chromosome Haplogroup A in Africa.]]
[[File:B haplogroup of Y-DNA.png|thumb|Distribution of haplogroup B (M60) of the human Y chromosome in native populations.]]
Various [[Y chromosome]] studies show that the San carry some of the most divergent (oldest) [[Human Y-chromosome DNA haplogroup|human Y-chromosome haplogroup]]s. These haplogroups are specific sub-groups of haplogroups [[Haplogroup A (Y-DNA)|A]] and [[Haplogroup B (Y-DNA)|B]], the two earliest branches on the human Y-chromosome [[phylogenetic tree|tree]].<ref name=j1>{{cite journal | vauthors = Knight A, Underhill PA, Mortensen HM, Zhivotovsky LA, Lin AA, Henn BM, Louis D, Ruhlen M, Mountain JL | display-authors = 6 | title = African Y chromosome and mtDNA divergence provides insight into the history of click languages | journal = Current Biology | volume = 13 | issue = 6 | pages = 464–473 | date = March 2003 | pmid = 12646128 | doi = 10.1016/S0960-9822(03)00130-1 | s2cid = 52862939 | doi-access = free }}</ref><ref>{{cite journal | vauthors = Hammer MF, Karafet TM, Redd AJ, Jarjanazi H, Santachiara-Benerecetti S, Soodyall H, Zegura SL | title = Hierarchical patterns of global human Y-chromosome diversity | journal = Molecular Biology and Evolution | volume = 18 | issue = 7 | pages = 1189–1203 | date = July 2001 | pmid = 11420360 | doi = 10.1093/oxfordjournals.molbev.a003906 | doi-access = free }}</ref><ref>{{cite journal | vauthors = Naidoo T, Schlebusch CM, Makkan H, Patel P, Mahabeer R, Erasmus JC, Soodyall H | title = Development of a single base extension method to resolve Y chromosome haplogroups in sub-Saharan African populations | journal = Investigative Genetics | volume = 1 | issue = 1 | pages = 6 | date = September 2010 | pmid = 21092339 | pmc = 2988483 | doi = 10.1186/2041-2223-1-6 }}</ref>
====Mitochondrial DNA====
[[Mitochondrial DNA]] studies also provide evidence that the San carry high frequencies of the earliest [[Human mitochondrial DNA haplogroup|haplogroup]] branches in the human mitochondrial DNA tree. This DNA is inherited only from one's mother. The most divergent (oldest) mitochondrial haplogroup, [[Haplogroup L0 (mtDNA)|L0]]d, has been identified at its highest frequencies in the southern African San groups.<ref name=j1/><ref>{{cite journal | vauthors = Chen YS, Olckers A, Schurr TG, Kogelnik AM, Huoponen K, Wallace DC | title = mtDNA variation in the South African Kung and Khwe-and their genetic relationships to other African populations | journal = American Journal of Human Genetics | volume = 66 | issue = 4 | pages = 1362–1383 | date = April 2000 | pmid = 10739760 | pmc = 1288201 | doi = 10.1086/302848 }}</ref><ref>{{cite journal | vauthors = Tishkoff SA, Gonder MK, Henn BM, Mortensen H, Knight A, Gignoux C, Fernandopulle N, Lema G, Nyambo TB, Ramakrishnan U, Reed FA, Mountain JL | display-authors = 6 | title = History of click-speaking populations of Africa inferred from mtDNA and Y chromosome genetic variation | journal = Molecular Biology and Evolution | volume = 24 | issue = 10 | pages = 2180–2195 | date = October 2007 | pmid = 17656633 | doi = 10.1093/molbev/msm155 | doi-access = free }}</ref><ref>{{cite journal | vauthors = Schlebusch CM, Naidoo T, Soodyall H | title = SNaPshot minisequencing to resolve mitochondrial macro-haplogroups found in Africa | journal = Electrophoresis | volume = 30 | issue = 21 | pages = 3657–3664 | date = November 2009 | pmid = 19810027 | doi = 10.1002/elps.200900197 | s2cid = 19515426 }}</ref>
====Autosomal DNA====
In a study published in March 2011, Brenna Henn and colleagues found that the ǂKhomani San, as well as the [[Sandawe people|Sandawe]] and [[Hadza people]]s of [[Tanzania]], were the most genetically diverse of any living humans studied. This high degree of genetic diversity hints at the origin of [[anatomically modern humans]].<ref>{{cite journal | vauthors = Henn BM, Gignoux CR, Jobin M, Granka JM, Macpherson JM, Kidd JM, Rodríguez-Botigué L, Ramachandran S, Hon L, Brisbin A, Lin AA, Underhill PA, Comas D, Kidd KK, Norman PJ, Parham P, Bustamante CD, Mountain JL, Feldman MW | display-authors = 6 | title = Hunter-gatherer genomic diversity suggests a southern African origin for modern humans | journal = Proceedings of the National Academy of Sciences of the United States of America | volume = 108 | issue = 13 | pages = 5154–5162 | date = March 2011 | pmid = 21383195 | pmc = 3069156 | doi = 10.1073/pnas.1017511108 | publisher = [[National Academy of Sciences]] | doi-access = free }}</ref><ref>{{cite journal |last=Kaplan |first=Matt |year=2011 |title=Gene Study Challenges Human Origins in Eastern Africa |journal=[[Scientific American]]|publisher=[[Nature Publishing Group]] |url=http://www.scientificamerican.com/article/gene-study-challenges-human-origin-africa/ |access-date=22 June 2012}}</ref>
====Medical DNA====
Among the ancient DNA from three [[hunter-gatherers]] sharing genetic similarity with [[San people]] and four Iron Age [[agriculturalists]], their [[SNPs]] indicated that they bore variants for resistance against [[sleeping sickness]] and [[Plasmodium vivax]].<ref name="Pfeiffer">{{cite journal | vauthors = Pfeiffer S | title = Disease as a Factor in the African Archaeological Record | journal = The African Archaeological Review | volume = 37 | issue = 3 | pages = 487–490 | year = 2020 | pmid = 32863518 | pmc = 7445818 | doi = 10.1007/s10437-020-09405-7 }}</ref> In particular, two out of the four Iron Age agriculturalists bore variants for resistance against sleeping sickness and three out of the four Iron Age agriculturalists bore [[Human genetic resistance to malaria#Duffy antigen receptor negativity|Duffy negative variants]] for resistance against [[malaria]].<ref name="Pfeiffer" /> In contrast to the Iron Age agriculturalists, from among the San-related hunter-gatherers, a six-year-old boy may have died from [[schistosomiasis]].<ref name="Pfeiffer" /> In [[Botswana]], a man, who dates to 1400 BP, may have also carried the Duffy negative variant for resistance against malaria.<ref name="Pfeiffer" />
==Recent African origin of modern humans==
{{Main|Recent African origin of modern humans}}
{{Further|Early human migrations|Basal Eurasian|Human genetic variation}}
Between 500,000 BP and 300,000 BP, [[anatomically modern humans]] may have emerged in Africa.<ref name="Pereira">{{cite journal | vauthors = Pereira L, Mutesa L, Tindana P, Ramsay M | title = African genetic diversity and adaptation inform a precision medicine agenda | journal = Nature Reviews. Genetics | volume = 22 | issue = 5 | pages = 284–306 | date = May 2021 | pmid = 33432191 | doi = 10.1038/s41576-020-00306-8 | publisher = Nature Reviews | s2cid = 231587564 }}</ref> As Africans (e.g., [[Y-Chromosomal Adam]], [[Mitochondrial Eve]]) have migrated from their places of origin in Africa to other locations in Africa, and as the time of divergence for [[East Africa]]n, [[Central Africa]]n, and [[West African]] lineages are similar to the time of divergence for the [[Southern African]] lineage, there is insufficient evidence to identify a specific region for the origin of humans in [[Africa]].<ref name="Bergström" /> In 100,000 BP, anatomically modern humans migrated from [[Africa]] into [[Eurasia]].<ref name="Benton">{{cite journal | vauthors = Benton ML, Abraham A, LaBella AL, Abbot P, Rokas A, Capra JA | title = The influence of evolutionary history on human health and disease | journal = Nature Reviews. Genetics | volume = 22 | issue = 5 | pages = 269–283 | date = May 2021 | pmid = 33408383 | pmc = 7787134 | doi = 10.1038/s41576-020-00305-9 }}</ref> Subsequently, tens of thousands of years after, the ancestors of all present-day Eurasians migrated from Africa into Eurasia and eventually became admixed with [[Denisovans]] and [[Neanderthals]].<ref name="Benton" />
==African historiography==
{{Main|African historiography}}
===Genetics===
Keita (2022) stated:<ref name="Keita">{{cite book |last1=Keita |first1=Shomarka O. |title=Studying Africa and Africans Today |date=2 September 2022 |publisher=B P International |pages=151-152 |chapter-url=https://www.researchgate.net/profile/Augustin-Holl/publication/363338424_Burial_Protocols_Megaliths_Production_and_Ancestor-Hood_in_Ancient_Senegambia_ca_1450_BCE_-_1500_CE/links/63745590431b1f5300a1516b/Burial-Protocols-Megaliths-Production-and-Ancestor-Hood-in-Ancient-Senegambia-ca-1450-BCE-1500-CE.pdf#page=158 |chapter=Genetics and African Historiography: Limitations and Insights |doi=10.9734/bpi/mono/978-93-5547-847-4/CH10 |isbn=978-93-5547-848-1}}</ref>
<blockquote>Historical questions addressed by geneticists have included population histories which focus on ancestral identities, migrations an[d] admixture at the individual and populations levels, with populations being defined or described in various ways, leading to units which are not really equivalent. Still other studies are more descriptive, provide basic information, and catalogue the variation in groups where there has been known interaction. None of this kind of work is without problems. [[Sampling (statistics)|Sample size]] and representativeness will likely for some time to come be a problem in such studies, even if old paradigms and ideas are successfully avoided—which requires a strong knowledge of the history of ideas. [[Statistical analysis|Statistical analyses]], heavily relied upon in much of this work, are not without problems and different results are possible depending on [[Statistical inference#Degree of models/assumptions|initial assumptions]], sample size, sample representativeness, and [[Statistics#Methods|methods]]. [[Sampling bias|Biases]] can be built into algorithms. Non-macro-evolutionary genetic history/historical genetics as in some sense a "new" kind of history has not yet been fully subjected to the kinds of philosophical treatment given on the one hand to paleontology, and the other hand to text based history—where scholars debate how much the past can really be known in some complete sense and attempt to identify the effect of [[Bias (statistics)|bias]] in description, [[Bias (statistics)#Interpretation|interpretation]] and explanation (McCullagh 2000). There is not yet a claimed or identifiable well-developed [[historiography]], in the sense of [[methodology]] of historical genetics or "genetic history" work which ironically, is not often published in [[List of history journals#Africa|history journals]] (de Chadeverian 2010, Egorova 2010).<ref name="Keita" /></blockquote>
Keita (2022) further stated: "The interest in genetics is often embedded in ideas about identity, which is a multilayered construct. Many populations will have a diversity of lineages as indicated by [[Y chromosome|male]] or [[Mitochondrial DNA#Female inheritance|female]] lineage DNA. Individuals may have varying ancestries which may not match their current social or ethnic identities. It is important to remember that a culturally defined group could [[Language shift|change language]] and gain different ancestry over time, unless social rules deny group inclusion to the descendants of such mating. A biologically defined group can change culture or languages. Languages can and have been adopted by new populations. These permutations depend on social or sociopolitical rules."<ref name="Keita" />
Yéré et al. (2022) stated: "The histories, knowledge and experiences of people in the African continent must drive scientific work that purports to be to their benefit. Leadership by scholars in Africa to discuss and unpack the kinds of assumptions that are made about race and ethnicity in African genomics is essential, as is an African research agenda that is interdisciplinary and attuned to the politics of biological characterization that is taking place in African genomics studies, where scholars from social and political science, history and ethics sit in the same spaces as those from biology and genomics. Only then can the history of African identity be better reflected in the concept of African genomes, and its hopes and limitations better understood."<ref name="Yéré">{{cite journal |last1=Yéré |first1=Henri-Michel |last2=Machirori |first2=Mavis |last3=De Vries |first3=Jantina |title=Unpacking race and ethnicity in African genomics research |journal=Nature Reviews Genetics |date=June 2022 |volume=23 |page=456 |doi=10.1038/s41576-022-00506-4 |pmid=35688877 |url=https://www.researchgate.net/profile/Henri-Yere/publication/361230166_Unpacking_race_and_ethnicity_in_African_genomics_research/links/62a888ba6886635d5cd9161b/Unpacking-race-and-ethnicity-in-African-genomics-research.pdf |issn=1471-0056 |oclc=9588452770 |s2cid=249574351}}</ref>
===Linguistics===
Güldemann (2018) stated:<ref name="Güldemann">{{cite book |last1=Güldemann |first1=Tom |title=The Languages and Linguistics of Africa |date=September 10, 2018 |publisher=De Gruyter Mouton |isbn=9783110421668 |pages=359–360 |chapter-url=https://pages.sandpoints.org/dotawo/library/Tom%20Guldemann/Historical%20Linguistics%20and%20Genealogical%20Language%20Classification%20in%20Africa%20(186)/Historical%20Linguistics%20and%20Genealogical%20La%20-%20Tom%20Guldemann.pdf |chapter=Historical linguistics and genealogical language classification in Africa |doi=10.1515/9783110421668-002 |oclc=1086052368 |s2cid=133888593}}</ref>
<blockquote>The reliance on Greenberg-like [[Joseph Greenberg#Languages of Africa|genealogical language classifications in Africa]] has had and still has important negative repercussions outside linguistics, especially in the disciplines concerned with human history like archaeology, genetics, etc. Flight (1981: 52) once wrote: "From a different point of view – for historians and prehistorians – the significance of [[Joseph Greenberg|Greenberg]]'s classification is no less obvious. The historical implications are immediate. A genetic classification of [[African languages]] is an outline plan for [[African history]]." It comes as no surprise that broad strokes of early African [[population history]], for example, by Heine (1979), MacDonald (1998), Ehret (1998, 2002), Blench (1999b, 2006a), etc. rely to a considerable extent on Greenberg’s classification, arguably misguiding basic assumptions about the history of Africa and its [[Demographics of Africa|peoples]]. An inspection of the literature makes clear that such a perception of Africa is even influential on the global level. To mention just an extreme example, Manning (2006: 139–141) speculates about the origin of most tropical language families in the Old World by practically deriving them from the equivocal [[Nilo-Saharan]] grouping in Africa.<ref name="Güldemann" /></blockquote>
== See also ==
*[[Genetic history of the African diaspora]]
== Notes ==
{{NoteFoot}}
== References ==
{{reflist}}
{{Human genetics}}
[[Category:Genetic genealogy]]
[[Category:History of Africa]]
[[Category:Modern human genetic history]]' |
