Editing Fula people (section)

==Genetics==
{{anchor|Genetics}}
The Fulani people are genetically an admixture of West and East African ancestries, specifically Niger-Congo and Nilo-Saharan components, but also display varying degrees of [[Eurasia|West Eurasian]] admixture through contact with groups from [[North Africa]].<ref>{{Cite journal |last1=Černý |first1=Viktor |last2=Fortes-Lima |first2=Cesar |last3=Tříska |first3=Petr |date=2021-04-26 |title=Demographic history and admixture dynamics in African Sahelian populations |journal=Human Molecular Genetics |volume=30 |issue=R1 |pages=R29–R36 |doi=10.1093/hmg/ddaa239 |issn=1460-2083 |pmid=33105478|doi-access=free }}</ref> The Fulani are the most wide-spread pastoralist group in the Sahel/Savannah belt.<ref name="doi.org">{{Cite journal |last1=Vicente |first1=Mário |last2=Priehodová |first2=Edita |last3=Diallo |first3=Issa |last4=Podgorná |first4=Eliška |last5=Poloni |first5=Estella S. |last6=Černý |first6=Viktor |last7=Schlebusch |first7=Carina M. |date=2019-12-02 |title=Population history and genetic adaptation of the Fulani nomads: inferences from genome-wide data and the lactase persistence trait |journal=BMC Genomics |volume=20 |issue=1 |pages=915 |doi=10.1186/s12864-019-6296-7 |issn=1471-2164 |pmc=6888939 |pmid=31791255 |doi-access=free }}</ref>

=== Paternal lineages (Y-DNA) ===
The paternal lineages of the Fula/Fulɓe/Fulani tend to vary depending on geographic location. According to a study by Cruciani et al. (2002), around 90% of Fulani individuals from [[Burkina Faso]] carried haplotype 24, which corresponds with the [[Haplogroup E-M2|E-M2]] (E1b1a) that is common in West Africa. The remainder belonged to haplotype 42/[[haplogroup E-M132]]. Both of these clades are today most frequent among [[Niger–Congo languages|Niger–Congo-speaking]] populations, particularly those inhabiting [[Senegal]]. Similarly, 53% of the Fulani in northern [[Cameroon]] bore haplogroup E-M132, with the rest mainly carrying other African clades (12% [[Macro-haplogroup A(X-BT)|haplogroup A]] and 6% haplogroup E1b1a). A significant minority carried the West Eurasian haplogroups [[Haplogroup T-M184|T]] (18%) and [[Haplogroup R1|R1]] (12%), making up together around ~30% of the total haplogroup variation.<ref name="Cruciani2002">{{cite journal | last1 = Cruciani | first1 = Fulvio | last2 = Santolamazza | first2 = Piero | last3 = Shen | first3 = Peidong | title = A Back Migration from Asia to 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 | doi=10.1086/340257 | pmid=11910562 | pmc=447595|display-authors=etal | date=May 2002}}</ref> Mulcare et al. (2004) observed a similar frequency of haplogroup R1 subclades in their Fulani samples from Cameroon (18%).<ref name="Mulcare">{{cite journal|title=The T Allele of a Single-Nucleotide Polymorphism 13.9 kb Upstream of the Lactase Gene (LCT) (C−13.9kbT) Does Not Predict or Cause the Lactase-Persistence Phenotype in Africans |year=2004 |last1=Mulcare |first1=Charlotte A. |last2=Weale |first2=Michael E. |last3=Jones |first3=Abigail L. |last4=Connell |first4=Bruce |last5=Zeitlyn |first5=David |last6=Tarekegn |first6=Ayele |last7=Swallow |first7=Dallas M. |last8=Bradman |first8=Neil |last9=Thomas |first9=Mark G. |journal=The American Journal of Human Genetics |volume=74 |issue=6 |pages=1102–1110 |doi=10.1086/421050 |pmid=15106124 |pmc=1182074}}</ref>

A study by Hassan et al. (2008) on a Fulani subgroup in [[Sudan]] observed a significantly higher occurrence of the West-Eurasian haplogroup R1 (53.8%). The remainder belonged to [[Haplogroup E-M215 (Y-DNA)|E-M215]] subclades, including 34.62% [[Haplogroup E-V68|E-M78]] and 27.2% [[Haplogroup E-V68#E-V22|E-V22]].<ref name="DoiajpaMissing">{{cite journal |doi=10.1002/ajpa.20876 |title=Y-chromosome variation among Sudanese: Restricted gene flow, concordance with language, geography, and history |year=2008 |last1=Hassan |first1=Hisham Y. |last2=Underhill |first2=Peter A. |last3=Cavalli-Sforza |first3=Luca L. |last4=Ibrahim |first4=Muntaser E. |journal=American Journal of Physical Anthropology |volume=137 |issue=3 |pages=316–23 |pmid=18618658}}</ref> Bučková et al. (2013) analyzed various Fulani subgroups, and observed [[Haplogroup R1b#R1b1b (R-V88)|R1b]] among the Fulani Zinder grouping with a frequency of ~31%. This was in sharp contrast to most of the other Fulani pastoralist groups elsewhere, including those from Burkina Faso, Cameroon, [[Mali]] and [[Chad]], which instead had nearly exclusive West African paternal haplogroups.<ref name="Bučková">{{cite journal|title=Multiple and differentiated contributions to the male gene pool of pastoral and farmer populations of the African Sahel |year=2013 |last1=Bučková |first1=Jana |last2=Cerný |first2=Viktor |last3=Novelletto |first3=Andrea |journal=American Journal of Physical Anthropology |volume=151 |pages=10–21 |pmid=23460272 |issue=1 |doi=10.1002/ajpa.22236 }}</ref>

===Maternal lineages (mtDNA)===
{| class="wikitable sortable"
|+ mtDNA Haplogroups of Fula Groups <ref>Černý, V., Kulichová, I., Poloni, E. S., Nunes, J. M., Pereira, L., Mayor, A., & Sanchez-Mazas, A. (2018). Genetic history of the African Sahelian populations. ''HLA'', 91(3), 153–166. https://doi.org/10.1111/tan.13189</ref>
|-
! Population !! African Sub-Saharan mtDNA !! Eurasian mtDNA (%)
|-
| Fulani Abalak (Niger) || 70% || 30%
|-
| Fulani Ader (Niger) || 80% || 20%
|-
| Fulani Balatungur (Niger) || 91% || 9%
|-
| Fulani Banfora (Burkina Faso) || 76% || 24%
|-
| Fulani Bongor (Chad) || 90% || 10%
|-
| Fulani Diffa (Niger) || 90% || 10%
|-
| Fulani Diafarabe (Mali) || 74% || 26%
|-
| Fulani Fouta Djallon (Guinea) || 83% || 17%
|-
| Fulani Ferlo (Senegal) || 98% || 2%
|-
| Fulani Linia (Chad) || 90% || 10%
|-
| Fulani Tcheboua (Cameroon) || 86% || 14%
|-
| Fulani Tindangou (Burkina Faso) || 80% || 20%
|-
| Fulani Zinder (Niger) || 90% || 10%
|-
| Fulani Ziniare (Burkina Faso) || 90% || 10%
|}
A study of four Fulani nomad populations (n = 186) in three Sahelian countries (Chad, Cameroon, and Burkina Faso), found that the only group of nomadic Fulani that manifests some similarities with geographically related agricultural populations (from Guinea-Bissau and Nigeria) comes from Tcheboua in northern Cameroon.<ref>{{Cite journal |last1=Cerný |first1=V. |last2=Hájek |first2=M. |last3=Bromová |first3=M. |last4=Cmejla |first4=R. |last5=Diallo |first5=I. |last6=Brdicka |first6=R. |date=2006 |title=MtDNA of Fulani nomads and their genetic relationships to neighboring sedentary populations |url=https://pubmed.ncbi.nlm.nih.gov/16900879/ |journal=Human Biology |volume=78 |issue=1 |pages=9–27 |doi=10.1353/hub.2006.0024 |issn=0018-7143 |pmid=16900879|s2cid=7045459 }}</ref><ref>{{Cite journal |last1=ČERNÝ |first1=V. |last2=HÁJEK |first2=M. |last3=BROMOVÁ |first3=M. |last4=ČMEJLA |first4=R. |last5=DIALLO |first5=I. |last6=BRDIČKA |first6=R. |date=2006 |title=mtDNA of Fulani Nomads and Their Genetic Relationships to Neighboring Sedentary Populations |url=https://www.jstor.org/stable/41466384 |journal=Human Biology |volume=78 |issue=1 |pages=9–27 |doi=10.1353/hub.2006.0024 |jstor=41466384 |pmid=16900879 |s2cid=7045459 |issn=0018-7143}}</ref>

===Autosomal DNA (overall)===
According to Tishkoff et al. (2009), the Fulani's genomic ancestry clusters near that of [[Chadic languages|Chadic]] and [[Central Sudanic languages|Central Sudanic]] speaking populations, with genetic affinities observed to the [[Hausa people]]. Based on this, the researchers suggest that the Fulani may have adopted a Niger-Congo language at some point in their history, while intermarrying with local populations. Additionally, moderate levels of West Eurasian admixture was also observed among the Fulani samples, which the authors propose may have been introduced via the [[Iberian Peninsula]] and Northern Africa.<ref name="Tishkoff">{{cite journal |title=The Genetic Structure and History of Africans and African Americans |year=2009 |last1=Tishkoff |first1=S. A. |last2=Reed |first2=F. A. |last3=Friedlaender |first3=F. R. |last4=Ehret |first4=C. |last5=Ranciaro |first5=A. |last6=Froment |first6=A. |last7=Hirbo |first7=J. B. |last8=Awomoyi |first8=A. A. |last9=Bodo |first9=J.-M. |last10=Doumbo |first10=O. |last11=Ibrahim |first11=M. |last12=Juma |first12=A. T. |last13=Kotze |first13=M. J. |last14=Lema |first14=G. |last15=Moore |first15=J. H. |last16=Mortensen |first16=H. |last17=Nyambo |first17=T. B. |last18=Omar |first18=S. A. |last19=Powell |first19=K. |last20=Pretorius |first20=G. S. |last21=Smith |first21=M. W. |last22=Thera |first22=M. A. |last23=Wambebe |first23=C. |last24=Weber |first24=J. L. |last25=Williams |first25=S. M. |journal=Science |volume=324 |issue=5930 |pages=1035–44 |pmid=19407144 |pmc=2947357 |doi=10.1126/science.1172257|bibcode=2009Sci...324.1035T }}</ref> Dobon et al. (2015), found that the Sudanese Fulani have largely ancestry from Niger-Kordofanian and Nilo-Saharan (Sudanic) speaking groups, with lower amounts of West-Eurasian ancestry.<ref>{{Cite journal |last1=Dobon |first1=Begoña |last2=Hassan |first2=Hisham Y. |last3=Laayouni |first3=Hafid |last4=Luisi |first4=Pierre |last5=Ricaño-Ponce |first5=Isis |last6=Zhernakova |first6=Alexandra |last7=Wijmenga |first7=Cisca |last8=Tahir |first8=Hanan |last9=Comas |first9=David |last10=Netea |first10=Mihai G. |last11=Bertranpetit |first11=Jaume |date=2015-05-28 |title=The genetics of East African populations: a Nilo-Saharan component in the African genetic landscape |journal=Scientific Reports |language=en |volume=5 |issue=1 |pages=9996 |doi=10.1038/srep09996 |pmid=26017457 |pmc=4446898 |bibcode=2015NatSR...5E9996D |issn=2045-2322}}</ref>
[[File:Woodabe young men.jpg|thumb|281x281px|Young Fulani men at [[Cure Salee]] festival, Niger.]]
Triska, Petr et al. (2015) showed that there is extensive admixture across the Sahel Belt, with the Fula carrying West African and East African components, as well as a Mozabite/North African component. These results support the hypothesis of a North African origin and a Western to Central Africa past migration for Fulani.<ref>{{Cite journal |last1=Triska |first1=Petr |last2=Soares |first2=Pedro |last3=Patin |first3=Etienne |last4=Fernandes |first4=Veronica |last5=Cerny |first5=Viktor |last6=Pereira |first6=Luisa |date=2015-11-26 |title=Extensive Admixture and Selective Pressure Across the Sahel Belt |journal=Genome Biology and Evolution |volume=7 |issue=12 |pages=3484–3495 |doi=10.1093/gbe/evv236 |issn=1759-6653 |pmc=4700964 |pmid=26614524}}</ref>

A full genome analysis was conducted by Vicente et al. in 2019, analyzing several different Fulani subgroups from various geographic regions. They found that the Fulani people are characterized by the admixture of local West African and East African components, but also display West-Eurasian admixture, mediated through historical North African groups. The West-Eurasian ancestry among Fulani was estimated to a mean average of 21,4% among the 53 samples from [[Ziniaré]] in [[Burkina Faso]].  According to the authors, there were two admixture events, the first being about 2000 years ago, with the second being more recent at around 300 years ago. This Eurasian ancestry was observed in the ancestry components of [[Mozabite people]]. They found that: "''Our findings suggest that Eurasian admixture and the European LP allele was introduced into the Fulani through contact with a North African population/s. We furthermore confirm the link between the lactose digestion phenotype in the Fulani to the MCM6/LCT locus by reporting the first GWAS of the lactase persistence trait. e observed a T-13910 allele frequency of 48.0%, while the genome-wide European admixture fraction in the Fulani is 21.4% at K = 3. The notable European admixture fraction in the Fulani coupled with the high frequencies of the LP T-13910 allele suggests the possibility of adaptive gene flow into the Fulani gene pool''".<ref name="doi.org"/> Another study in 2020 by Priehodová et al., suggest an older date for the introduction of one variant of the LP allele in the Sahel, about ~8.5 ka.<ref>{{Cite journal |last1=Priehodová |first1=Edita |last2=Austerlitz |first2=Frédéric |last3=Čížková |first3=Martina |last4=Nováčková |first4=Jana |last5=Ricaut |first5=François-Xavier |last6=Hofmanová |first6=Zuzana |last7=Schlebusch |first7=Carina M. |last8=Černý |first8=Viktor |date=November 2020 |title=Sahelian pastoralism from the perspective of variants associated with lactase persistence |url=https://pubmed.ncbi.nlm.nih.gov/32812238/ |journal=American Journal of Physical Anthropology |volume=173 |issue=3 |pages=423–436 |doi=10.1002/ajpa.24116 |issn=1096-8644 |pmid=32812238|s2cid=221179656 }}</ref>

A study in 2019 by Fan et al., found that the Fulani sampled from Cameroon, clustered with Afro-Asiatic speakers from East Africa in the phylogenetic analysis, which the authors said indicates a potential shift in language to Niger-Congo. The analysis on autosomal markers found traces of West Eurasian-related ancestry in this population, which suggests a North African or East African origin (as North and East Africans also have such ancestry likely related to expansions of farmers and herders from the Near East) and is consistent with the presence at moderate frequency of the −13,910T variant associated with lactose tolerance in European populations.<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 |journal=Genome Biology |volume=20 |issue=1 |pages=82 |doi=10.1186/s13059-019-1679-2 |issn=1474-760X |pmc=6485071 |pmid=31023338 |doi-access=free }}</ref>

In 2020, a study inferred that the Fulani of western Cameroon have 48% Mende-related, 23% East African-related, and 29% non-African-related ancestry.<ref>{{Cite journal |last1=Lipson |first1=Mark |last2=Ribot |first2=Isabelle |last3=Mallick |first3=Swapan |last4=Rohland |first4=Nadin |last5=Olalde |first5=Iñigo |last6=Adamski |first6=Nicole |last7=Broomandkhoshbacht |first7=Nasreen |last8=Lawson |first8=Ann Marie |last9=López |first9=Saioa |last10=Oppenheimer |first10=Jonas |last11=Stewardson |first11=Kristin |last12=Asombang |first12=Raymond Neba’ane |last13=Bocherens |first13=Hervé |last14=Bradman |first14=Neil |last15=Culleton |first15=Brendan J. |date=2020-01-22 |title=Ancient West African foragers in the context of African population history |url=https://reich.hms.harvard.edu/sites/reich.hms.harvard.edu/files/inline-files/Shum_Laka_SI.pdf |journal=Nature |volume=577 |issue=7792 |pages=665–670 |bibcode=2020Natur.577..665L |doi=10.1038/s41586-020-1929-1 |issn=0028-0836 |pmc=8386425 |pmid=31969706}}</ref>

In 2023, whole genomes of Fulani individuals from various Sahelian samples were analyzed, and the researches said the non-Sub-Saharan genetic ancestry within the Fulani cannot be solely explained by recent admixture events. Fulani may be descendants of Saharan cattle herders during the last Green Sahara, who had some genomic similarities to Late Neolithic Moroccans based on ancient samples.<ref>{{Cite journal |last=D’Atanasio |first=Eugenia |date=April 6, 2023 |title=Echoes from the last Green Sahara: whole genome analysis of Fulani, a key population to unveil the genetic evolutionary history of Africa |url=https://www.biorxiv.org/content/10.1101/2023.04.06.535569v1 |website=bioRxiv|doi=10.1101/2023.04.06.535569 |s2cid=258041998 }}</ref>

Another 2023 study inferred that "The Fulani derived 50% of their ancestry from a population related to the [[Amhara people|Amhara]] and 50% from a population related to the [[Tikar people|Tikari]] (consistent with TreeMix results with 3 migration events)."<ref>{{Cite journal |last1=Fan |first1=Shaohua |last2=Spence |first2=Jeffrey P. |last3=Feng |first3=Yuanqing |last4=Hansen |first4=Matthew E. B. |last5=Terhorst |first5=Jonathan |last6=Beltrame |first6=Marcia H. |last7=Ranciaro |first7=Alessia |last8=Hirbo |first8=Jibril |last9=Beggs |first9=William |last10=Thomas |first10=Neil |last11=Nyambo |first11=Thomas |last12=Mpoloka |first12=Sununguko Wata |last13=Mokone |first13=Gaonyadiwe George |last14=Njamnshi |first14=Alfred |last15=Folkunang |first15=Charles |date=2023-03-02 |title=Whole-genome sequencing reveals a complex African population demographic history and signatures of local adaptation |journal=Cell |volume=186 |issue=5 |pages=923–939.e14 |doi=10.1016/j.cell.2023.01.042 |issn=1097-4172 |pmid=36868214|pmc=10568978 }}</ref>