This review focuses on the Sahel/Savannah belt, a large region of Africa where two alternative subsistence systems (pastoralism and agriculture), nowadays, interact. It is a long-standing question whether the pastoralists became isolated here from other populations after cattle began to spread into Africa (~8 thousand years ago, kya) or, rather, began to merge with other populations, such as agropastoralists, after the domestication of sorghum and pearl millet (~5 kya) and with the subsequent spread of agriculture. If we look at lactase persistence, a trait closely associated with pastoral lifestyle, we see that its variants in current pastoralists distinguish them from their farmer neighbours. Most other (mostly neutral) genetic polymorphisms do not, however, indicate such clear differentiation between these groups; they suggest a common origin and/or an extensive gene flow. Genetic affinity and ecological symbiosis between the two subsistence systems can help us better understand the population history of this African region. In this review, we show that genomic datasets of modern Sahel/Savannah belt populations properly collected in local populations can complement the still insufficient archaeological research of this region, especially when dealing with the prehistory of mobile populations with perishable material culture and therefore precarious archaeological visibility.

Archaeogenetics Laboratory Institute of Archaeology of the Academy of Sciences of the Czech Republic Letenská 1 118 01 Prague Czech Republic

Human Evolution Department of Organismal Biology Evolutionary Biology Centre Uppsala University Norbyvägen 18C 752 36 Uppsala Sweden

Zobrazit více v PubMed

Raynaut C., Delville P.L. Un espace partagé: Complémentarité et concurrence des usages. In: Raynaut C., editor. Sahels Diversité et Dynamiques des Relations Sociétés-Nature. Karthala; Paris, France: 1997. pp. 143–174.

Pedersen J., Benjaminsen T.A. One Leg or Two? Food Security and Pastoralism in the Northern Sahel. Hum. Ecol. 2008;36:43–57. doi: 10.1007/s10745-007-9136-3. DOI

Hardin G. The tragedy of the commons. The population problem has no technical solution; it requires a fundamental extension in morality. Science. 1968;162:1243–1248. doi: 10.1126/science.162.3859.1243. PubMed DOI

Behnke R. Earth system sciences. In: Mortimore M., editor. The End of Desertification? Disputing Environmental Change in the Drylands. Springer; Berlin/Heidelberg, Germany: 2016.

Dupire M. Étude Descriptive des Wodaabe du Sahel Nigérien. Karthala; Paris, France: 1962. Peuls Nomads.

Stenning D.J. Savannah Nomads; A Study of the Wodaabe Pastoral Fulani of Western Bornu Province, Northern Region, Nigeria. International African Institute by Oxford University Press; London, UK: 1959.

Asad T. Power, Authority and Consent in a Nomadic Tribe. C Hurst and Company; New York, NY, USA: 1970. The Kababish Arabs.

MacMichael H.A. The Tribes of Northern and Central Kordofán. Frank Cass & Co., Ltd.; London, UK: 1967.

Baroin C. Anarchie et Cohésion Sociale chez les Toubou: Les Daza Késerda (Niger) Cambridge University Press; Cambridge, UK: Editions de la Maison des Sciences de L’homme; Paris, France: 1985.

Chapelle J. Nomades Noires du Sahara. Harmattan; Paris, France: 1967. Les Toubous.

Tubiana M.-J. De Trupeaux et de Femmes. Mariage et Transferts de Biens Chez les Beri (Zaghawa et Bideyat) du Tchad et du Soudan. Harmattan; Paris, France: 1985.

Lhote H. Les Touaregs du Hoggar (Ahaggar) Payot; Paris, France: 1955.

Rodd R. People of the Veil; Being an Account of the Habits, Organisation and History of the Wandering Tuareg Tribes Which Inhabit the Mountains of Air, or Asben in the Central Sahara. Macmillan; London, UK: 1926.

Casajus D. La Société et les Morts Chez les Touaregs Kel Ferwan. Cambridge University Press; Cambridge, UK: 1987. La tente dans la Solitude.

Xiao N., Cai S., Moritz M., Garabed R., Pomeroy L.W. Spatial and Temporal Characteristics of Pastoral Mobility in the Far North Region, Cameroon: Data Analysis and Modeling. PLoS ONE. 2015;10:e0131697. doi: 10.1371/journal.pone.0131697. PubMed DOI PMC

Prussin L. Afrucan Nomadic Architecture: Space, Place and Gender. Smithsonian; Washington, WA, USA: 1995.

Breman H., de Wit C.T. Rangeland productivity and exploitation in the sahel. Science. 1983;221:1341–1347. doi: 10.1126/science.221.4618.1341. PubMed DOI

Brierley C., Manning K., Maslin M. Pastoralism may have delayed the end of the green Sahara. Nat. Commun. 2018;9:4018. doi: 10.1038/s41467-018-06321-y. PubMed DOI PMC

Homewood K. Ecology of African Pastoralist Societies. James Currey; Oxford, UK: 2008.

Wendorf F., Schild R. Holocene Settlement of the Egyptian Sahara: Volume 1: The Archaeology of Nabta Playa. Kluwer Academic/Plenum Publishers; New York, NY, USA: 2001.

Wendorf F., Schild R., Close A.E. Cattle-Keepers of the Eastern Sahara. The Neolithic of Bir Kiseiba. Department of Anthropology and Institute for the Study of Earth and Man, Southern Methodist University; Dallas, TX, USA: 1984.

Dunne J., di Lernia S., Chłodnicki M., Kherbouche F., Evershed R. Timing and pace of dairying inception and animal husbandry practices across Holocene North Africa. Quat. Int. 2018;471:147–159. doi: 10.1016/j.quaint.2017.06.062. DOI

Neumann K. The romance of farming: Plant cultivation and domestication in Africa. In: Stahl A.B., editor. African Archaeology: A Critical Introduction. Blackwell Publishing Ltd.; Malden, MA, USA: 2005. pp. 249–275.

Phelps L.N., Broennimann O., Manning K., Timpson A., Jousse H., Mariethoz G., Fordham D.A., Shanahan T.M., Davis B.A.S., Guisan A. Reconstructing the climatic niche breadth of land use for animal production during the African Holocene. Glob. Ecol. Biogeogr. 2020;29:127–147. doi: 10.1111/geb.13015. DOI

Lhote H. The search for the Tassili frescoes. In: Lhote H., editor. The Story of the Prehistoric Rock-Paintings of the Sahara. Dutton; New York, NY, USA: 1959.

Le Quellec J.-L. Recent Work on Saharan rock art (2010–2014) In: Bahn P., Franklin N., Strecker M., Devlet E., editors. Rock Art Studies: News of the World V. Archaeopress Publishing; Oxford, UK: 2016. pp. 55–74.

Wright D.K. Humans as Agents in the Termination of the African Humid Period. Front. Earth Sci. 2017;5:4. doi: 10.3389/feart.2017.00004. DOI

Linseele V. From the first stock keepers to specialised pastoralists in the West African Savannah. In: Bollig M., Schnegg M., Wotzka H.-P., editors. Pastoralism in Africa: Past, Present and Future. Berghan Books; New York, NY, USA: 2013. pp. 145–170.

Sereno P.C., Garcea E.A., Jousse H., Stojanowski C.M., Saliege J.F., Maga A., Ide O.A., Knudson K.J., Mercuri A.M., Stafford T.W., Jr., et al. Lakeside cemeteries in the Sahara: 5000 years of holocene population and environmental change. PLoS ONE. 2008;3:e2995. doi: 10.1371/journal.pone.0002995. PubMed DOI PMC

Bâ A.H., Daget J., Monod T. L’empire Peul du Macina: (1818–1853) Editions de l’Ecole des Hautes Études en Sciences Sociales; Paris, France: 1984.

Shahack-Gross R. Implications for the Spread of Pastoralism in East Africa. Saint Louis University; Saint Louis, MS, USA: 2002. Taphonomy of abandoned Maasai settlememnts in southern Kenya with special reference to livestock enclosures.

Černý V., Pereira L., Musilová E., Kujanová M., Vašíková A., Blasi P., Garofalo L., Soares P., Diallo I., Brdička R., et al. Genetic structure of pastoral and farmer populations in the African Sahel. Mol. Biol. Evol. 2011;28:2491–2500. doi: 10.1093/molbev/msr067. PubMed DOI

Kulichová I., Fernandes V., Deme A., Nováčková J., Stenzl V., Novelletto A., Pereira L., Černý V. Internal diversification of non-Sub-Saharan haplogroups in Sahelian populations and the spread of pastoralism beyond the Sahara. Am. J. Phys. Anthropol. 2017;164:424–434. doi: 10.1002/ajpa.23285. PubMed DOI

Nováčková J., Čížková M., Mokhtar M.G., Duda P., Stenzl V., Tříska P., Hofmanová Z., Černý V. Subsistence strategy was the main factor driving population differentiation in the bidirectional corridor of the African Sahel. Am. J. Phys. Anthropol. 2020;171:496–508. doi: 10.1002/ajpa.24001. PubMed DOI

Ranciaro A., Campbell M.C., Hirbo J.B., Ko W.Y., Froment A., Anagnostou P., Kotze M.J., Ibrahim M., Nyambo T., Omar S.A., et al. Genetic origins of lactase persistence and the spread of pastoralism in Africa. Am. J. Hum. Genet. 2014;94:496–510. doi: 10.1016/j.ajhg.2014.02.009. PubMed DOI PMC

Ségurel L., Bon C. On the Evolution of Lactase Persistence in Humans. Annu. Rev. Genomics. Hum. Genet. 2017;18:297–319. doi: 10.1146/annurev-genom-091416-035340. PubMed DOI

Priehodová E., Austerlitz F., Čížková M., Nováčková J., Ricaut F.-X., Hofmanová Z., Schlebusch C.M., Černý V. Sahelian pastoralism from the perspective of variants associated with lactase persistence. Am. J. Phys. Anthropol. 2020;173:423–436. doi: 10.1002/ajpa.24116. PubMed DOI

Priehodová E., Austerlitz F., Čížková M., Mokhtar M.G., Poloni E.S., Černý V. The historical spread of Arabian Pastoralists to the eastern African Sahel evidenced by the lactase persistence −13,915*G allele and mitochondrial DNA. Am. J. Hum. Biol. 2017;29:e22950. doi: 10.1002/ajhb.22950. PubMed DOI

Priehodová E., Abdelsawy A., Heyer E., Černý V. Lactase Persistence Variants in Arabia and in the African Arabs. Hum. Biol. 2014;86:7–18. doi: 10.3378/027.086.0101. PubMed DOI

Černý V., Fortes-Lima C., Tříska P. Demographic history and admixture dynamics in African Sahelian populations. Hum. Mol. Genet. 2021;30:R29–R36. doi: 10.1093/hmg/ddaa239. PubMed DOI

Campbell M.C., Ranciaro A. Human adaptation, demography and cattle domestication: An overview of the complexity of lactase persistence in Africa. Hum. Mol. Genet. 2021;30:R98–R109. doi: 10.1093/hmg/ddab027. PubMed DOI

Vicente M., Priehodová E., Diallo I., Podgorná E., Poloni E.S., Černý V., Schlebusch C.M. Population history and genetic adaptation of the Fulani nomads: Inferences from genome-wide data and the lactase persistence trait. BMC Genom. 2019;20:915. doi: 10.1186/s12864-019-6296-7. PubMed DOI PMC

Enattah N.S., Jensen T.G., Nielsen M., Lewinski R., Kuokkanen M., Rasinpera H., El-Shanti H., Seo J.K., Alifrangis M., Khalil I.F., et al. Independent introduction of two lactase-persistence alleles into human populations reflects different history of adaptation to milk culture. Am. J. Hum. Genet. 2008;82:57–72. doi: 10.1016/j.ajhg.2007.09.012. PubMed DOI PMC

Černý V., Kulichová I., Poloni E.S., Nunes J.M., Pereira L., Mayor A., Sanchez-Mazas A. Genetic history of the African Sahelian populations. Hla. 2018;91:153–166. doi: 10.1111/tan.13189. PubMed DOI

Tříska P., Soares P., Patin E., Fernandes V., Černý V., Pereira L. Extensive admixture and selective pressure across the Sahel Belt. Genome Biol. Evol. 2015;7:484–3495. doi: 10.1093/gbe/evv236. PubMed DOI PMC

Sanchez-Mazas A., Černý V., Da D., Buhler S., Podgorná E., Chevallier E., Brunet L., Weber S., Kervaire B., Testi M., et al. The HLA-B landscape of Africa: Signatures of pathogen-driven selection and molecular identification of candidate alleles to malaria protection. Mol. Ecol. 2017;26:6238–6252. doi: 10.1111/mec.14366. PubMed DOI

Luoni G., Verra F., Arca B., Sirima B.S., Troye-Blomberg M., Coluzzi M., Kwiatkowski D., Modiano D. Antimalarial antibody levels and IL4 polymorphism in the Fulani of West Africa. Genes Immun. 2001;2:411–414. doi: 10.1038/sj.gene.6363797. PubMed DOI

Modiano D., Luoni G., Sirima B.S., Lanfrancotti A., Petrarca V., Cruciani F., Simpore J., Ciminelli B.M., Foglietta E., Grisanti P., et al. The lower susceptibility to Plasmodium falciparum malaria of Fulani of Burkina Faso (west Africa) is associated with low frequencies of classic malaria-resistance genes. Trans. R. Soc. Trop. Med. Hyg. 2001;95:149–152. doi: 10.1016/S0035-9203(01)90141-5. PubMed DOI

Boström S., Giusti P., Arama C., Persson J.O., Dara V., Traore B., Dolo A., Doumbo O., Troye-Blomberg M. Changes in the levels of cytokines, chemokines and malaria-specific antibodies in response to Plasmodium falciparum infection in children living in sympatry in Mali. Malar. J. 2012;11:109. doi: 10.1186/1475-2875-11-109. PubMed DOI PMC

Quin J.E., Bujila I., Cherif M., Sanou G.S., Qu Y., Vafa Homann M., Rolicka A., Sirima S.B., O’Connell M.A., Lennartsson A., et al. Major transcriptional changes observed in the Fulani, an ethnic group less susceptible to malaria. Elife. 2017;6:29156. doi: 10.7554/eLife.29156. PubMed DOI PMC

Heyer E., Chaix R., Pavard S., Austerlitz F. Sex-specific demographic behaviours that shape human genomic variation. Mol. Ecol. 2012;21:597–612. doi: 10.1111/j.1365-294X.2011.05406.x. PubMed DOI

Garcia-Fernandez C., Font-Porterias N., Kucinskas V., Sukarova-Stefanovska E., Pamjav H., Makukh H., Dobon B., Bertranpetit J., Netea M.G., Calafell F., et al. Sex-biased patterns shaped the genetic history of Roma. Sci. Rep. 2020;10:14464. doi: 10.1038/s41598-020-71066-y. PubMed DOI PMC

Silva M., Oliveira M., Vieira D., Brandao A., Rito T., Pereira J.B., Fraser R.M., Hudson B., Gandini F., Edwards C., et al. A genetic chronology for the Indian Subcontinent points to heavily sex-biased dispersals. BMC Evol. Biol. 2017;17:88. doi: 10.1186/s12862-017-0936-9. PubMed DOI PMC

Shriner D., Rotimi C.N. Genetic history of Chad. Am. J. Phys. Anthropol. 2018;167:804–812. doi: 10.1002/ajpa.23711. PubMed DOI PMC

Čížková M., Munclinger P., Diallo M.Y., Kulichová I., Mokhtar M.G., Dème A., Pereira L., Černý V. Genetic Structure of the Western and Eastern African Sahel/Savannah Belt and the Role of Nomadic Pastoralists as Inferred from the Variation of D-Loop Mitochondrial DNA Sequences. Hum. Biol. 2017;89:281–302. doi: 10.13110/humanbiology.89.4.02. PubMed DOI

Bryc K., Auton A., Nelson M.R., Oksenberg J.R., Hauser S.L., Williams S., Froment A., Bodo J.M., Wambebe C., Tishkoff SA et a.l. Genome-wide patterns of population structure and admixture in West Africans and African Americans. Proc. Natl. Acad. Sci. USA. 2010;107:786–791. doi: 10.1073/pnas.0909559107. PubMed DOI PMC

Korunes K.L., Soares-Souza G.B., Bobrek K., Tang H., Araújo I.I., Goldberg A., Beleza S. Sex-biased admixture and assortative mating shape genetic variation and influence demographic inference in admixed Cabo Verdeans. G3. 2022;12:jkac183. doi: 10.1093/g3journal/jkac183. PubMed DOI PMC

Fortes-Lima C., Gessain A., Ruiz-Linares A., Bortolini M.C., Migot-Nabias F., Bellis G., Moreno-Mayar J.V., Restrepo B.N., Rojas W., Avendano-Tamayo E., et al. Genome-wide Ancestry and Demographic History of African-Descendant Maroon Communities from French Guiana and Suriname. Am. J. Hum. Genet. 2017;101:725–736. doi: 10.1016/j.ajhg.2017.09.021. PubMed DOI PMC

Fortes-Lima C., Tříska P., Čížková M., Podgorná E., Diallo M.Y., Schlebusch C.M., Černý V. Demographic and Selection Histories of Populations Across the Sahel/Savannah Belt. Mol. Biol. Evol. 2022;39:msac209. doi: 10.1093/molbev/msac209. PubMed DOI PMC

Alexander D.H., Lange K. Enhancements to the ADMIXTURE algorithm for individual ancestry estimation. BMC Bioinform. 2011;12:246. doi: 10.1186/1471-2105-12-246. PubMed DOI PMC

Diaz-Papkovich A., Anderson-Trocme L., Ben-Eghan C., Gravel S. UMAP reveals cryptic population structure and phenotype heterogeneity in large genomic cohorts. PLoS Genet. 2019;15:e1008432. doi: 10.1371/journal.pgen.1008432. PubMed DOI PMC

Auton A., Brooks L.D., Durbin R.M., Garrison E.P., Kang H.M., Korbel J.O., Marchini J.L., McCarthy S., McVean G.A., Abecasis G.R. A global reference for human genetic variation. Nature. 2015;526:68–74. PubMed PMC

Gurdasani D., Carstensen T., Tekola-Ayele F., Pagani L., Tachmazidou I., Hatzikotoulas K., Karthikeyan S., Iles L., Pollard M.O., Choudhury A., et al. The African Genome Variation Project shapes medical genetics in Africa. Nature. 2015;517:327–332. doi: 10.1038/nature13997. PubMed DOI PMC

Haber M., Mezzavilla M., Bergstrom A., Prado-Martinez J., Hallast P., Saif-Ali R., Al-Habori M., Dedoussis G., Zeggini E., Blue-Smith J., et al. Chad Genetic Diversity Reveals an African History Marked by Multiple Holocene Eurasian Migrations. Am. J. Hum. Genet. 2016;99:1316–1324. doi: 10.1016/j.ajhg.2016.10.012. PubMed DOI PMC

Hollfelder N., Schlebusch C.M., Gunther T., Babiker H., Hassan H.Y., Jakobsson M. Northeast African genomic variation shaped by the continuity of indigenous groups and Eurasian migrations. PLoS Genet. 2017;13:e1006976. doi: 10.1371/journal.pgen.1006976. PubMed DOI PMC

Fernandes V., Brucato N., Ferreira J.C., Pedro N., Cavadas B., Ricaut F.X., Alshamali F., Pereira L. Genome-Wide Characterization of Arabian Peninsula Populations: Shedding Light on the History of a Fundamental Bridge between Continents. Mol. Biol. Evol. 2019;36:575–586. doi: 10.1093/molbev/msz005. PubMed DOI

Hernandez C.L., Pita G., Cavadas B., Lopez S., Sanchez-Martinez L.J., Dugoujon J.M., Novelletto A., Cuesta P., Pereira L., Calderon R. Human Genomic Diversity Where the Mediterranean Joins the Atlantic. Mol. Biol. Evol. 2020;37:1041–1055. doi: 10.1093/molbev/msz288. PubMed DOI PMC

Patterson N., Moorjani P., Luo Y., Mallick S., Rohland N., Zhan Y., Genschoreck T., Webster T., Reich D. Ancient admixture in human history. Genetics. 2012;192:1065–1093. doi: 10.1534/genetics.112.145037. PubMed DOI PMC

Diallo M.Y., Čížková M., Kulichová I., Podgorná E., Priehodová E., Nováčková J., Fernandes V., Pereira L., Černý V. Circum-Saharan Prehistory through the Lens of mtDNA Diversity. Genes. 2022;13:533. doi: 10.3390/genes13030533. PubMed DOI PMC

Young W.C. The Rashaayda Bedouin: Arab Pastoralists of Eastern Sudan. Harcourt Brace College Publishers; Fort Worth, UK: 1996.

Lipson M., Ribot I., Mallick S., Rohland N., Olalde I., Adamski N., Broomandkhoshbacht N., Lawson A.M., Lopez S., Oppenheimer J., et al. Ancient West African foragers in the context of African population history. Nature. 2020;577:665–670. doi: 10.1038/s41586-020-1929-1. PubMed DOI PMC

Fu Q., Hajdinjak M., Moldovan O.T., Constantin S., Mallick S., Skoglund P., Patterson N., Rohland N., Lazaridis I., Nickel B., et al. An early modern human from Romania with a recent Neanderthal ancestor. Nature. 2015;524:216–219. doi: 10.1038/nature14558. PubMed DOI PMC

Duckworth C.N., Cuénod A., Mattingly D.J. Mobile Technologies in the Ancient Sahara and Beyond. Cambridge University Press; Cambridge, UK: 2020.

Maley J., Vernet R. Populations and Climatic Evolution in North Tropical Africa from the End of the Neolithic to the Dawn of the Modern Era. Afr. Archaeol. Rev. 2015;32:179–232. doi: 10.1007/s10437-015-9190-y. DOI

Lovejoy P.E. Salt of the Desert Sun: A History of Salt Production and Trade in the Central Sudan. Cambridge University Press; Cambridge, UK: 1986.

McDougall E.A. Salts of the Western Sahara: Myths, Mysteries, and Historical Significance. Int. J. Afr. Hist. Stud. 1990;23:231–257. doi: 10.2307/219336. DOI

McIntosh S.K. Long-Distance Exchange and Urban Trajectories in the First Millennium AD. Case Studies from the Middle Niger and Middle Senegal River Valleys. In: Sterry M., Mattingly D.J., editors. Urbanisation and State Formation in the Ancient Sahara and Beyond. Cambridge University Press; Cambridge, UK: 2020. pp. 521–563.

Sherratt A. Plough and pastoralism: Aspects of the secondary products revolution. In: Hodder I., Isaac G., Hammond N., editors. Pattern of the Past: Studies in Honour of David Clarke. Cambridge University Press; Cambridge, UK: 1981. pp. 261–305.

Retsö J. The domestication of the camel and the establisment of the frankincence road from South Arabia. Orient. Suec. 1991;40:187–219.

Braukämper U. Notes on the origin of Baggara Arab culture with special reference to the Shuwa. In: Ownes J., editor. Arabs and Arabic in the Lake Chad Region. Volume Sprache und Geschichte in Africa 14. Rüdiger Köppe Verlag; Köln, German: 1994. pp. 13–46.

Schultz E. From Pagan to Pullo: Ethnic Identity Change in Northern Cameroon. Afr. J. Int. Afr. Inst. 1984;54:46–64. doi: 10.2307/1160143. DOI

Barth F. Ethnic groups and boundaries. The Social Organization of Culture Difference. Universitetsforlaget; Oslo, Norway: 1969.

Mohammadou E. Traditions Historiques des Peuples du Cameroun Central, Mbere et Mboum Tikar. Volume 1 Ilcaa; Tokyo, Japan: 1990.

de Haan C., Dubern E., Garancher B., Quintero C. Pastoralism Development in the Sahel A Road to Stability? International Bank for Reconstruction and Development; Washington, WA, USA: 2016.

Ickowicz A., Ancey V., Corniaux C., Duteurtre G., Poccard-Chappuis R., Touré I., Vall E., Wane A. Crop–livestock production systems in the Sahel–increasing resilience for adaptation to climate change and preserving food security. In: Meybeck A., Lankoski J., Redfern S., Azzu N., Gitz V., editors. Building Resilience for Adaptation to Climate Change in the Agriculture Sector. FAO; Rome, Italy: 2012.

Swift J. Sahelian Pastoralists: Underdevelopment, Desertification, and Famine. Annu. Rev. Anthropol. 1977;6:457–478. doi: 10.1146/annurev.an.06.100177.002325. DOI

Simonet C., Traoré S.M., Brunelin S., Royer L. Livestick Markets in the Sahel: Market Integration and the role of Climate and Cconflict in Price Formation. BRACED; London, UK: 2020.

Pereira L., Mutesa L., Tindana P., Ramsay M. African genetic diversity and adaptation inform a precision medicine agenda. Nat. Rev. Genet. 2021;22:284–306. doi: 10.1038/s41576-020-00306-8. PubMed DOI

Scheinfeldt L.B., Soi S., Tishkoff S.A. Colloquium paper: Working toward a synthesis of archaeological, linguistic, and genetic data for inferring African population history. Proc. Natl. Acad. Sci. USA. 2010;107((Suppl. 2)):8931–8938. doi: 10.1073/pnas.1002563107. PubMed DOI PMC

Hammer M.F., Woerner A.E., Mendez F.L., Watkins J.C., Wall J.D. Genetic evidence for archaic admixture in Africa. Proc. Natl. Acad. Sci. USA. 2011;108:15123–15128. doi: 10.1073/pnas.1109300108. PubMed DOI PMC

Xu D., Pavlidis P., Taskent R.O., Alachiotis N., Flanagan C., DeGiorgio M., Blekhman R., Ruhl S., Gokcumen O. Archaic Hominin Introgression in Africa Contributes to Functional Salivary MUC7 Genetic Variation. Mol. Biol. Evol. 2017;34:2704–2715. doi: 10.1093/molbev/msx206. PubMed DOI PMC

Population history and admixture of the Fulani people from the Sahel