Haplogroup L3 is a human mitochondrial DNA (mtDNA)

haplogroup A haplotype is a group of alleles in an organism that are inherited together from a single parent, and a haplogroup (haploid from the el, ἁπλοῦς, ''haploûs'', "onefold, simple" and en, group) is a group of similar haplotypes that share ...

. The clade has played a pivotal role in the early dispersal of

anatomically modern human Anatomy () is the branch of biology concerned with the study of the structure of organisms and their parts. Anatomy is a branch of natural science that deals with the structural organization of living things. It is an old science, having its ...

s. It is strongly associated with the out-of-Africa migration of modern humans of about 70–50,000 years ago. It is inherited by all modern non-African populations, as well as by some populations in Africa.

Origin

Haplogroup L3 arose close to 70,000 years ago, near the time of the recent out-of-Africa event. This dispersal originated in

East Africa East Africa, Eastern Africa, or East of Africa, is the eastern subregion of the African continent. In the United Nations Statistics Division scheme of geographic regions, 10-11-(16*) territories make up Eastern Africa: Due to the historica ...

and expanded to West Asia, and further to South and Southeast Asia in the course of a few millennia, and some research suggests that L3 participated in this migration out of Africa. A 2007 estimate for the age of L3 suggested a range of 104–84,000 years ago. More recent analyses, including Soares et al. (2012) arrive at a more recent date, of roughly 70–60,000 years ago. Soares et al. also suggest that L3 most likely expanded from East Africa into Eurasia sometime around 65–55,000 years ago years ago as part of the recent out-of-Africa event, as well as from East Africa into Central Africa from 60 to 35,000 years ago. In 2016, Soares et al. again suggested that haplogroup L3 emerged in East Africa, leading to the Out-of-Africa migration, around 70–60,000 years ago. Haplogroups L6 and L4 form sister clades of L3 which arose in East Africa at roughly the same time but which did not participate in the out-of-Africa migration. The ancestral clade L3'4'6 has been estimated at 110 kya, and the L3'4 clade at 95 kya. Africa origin thesis for L3 Fig4 HTML

The possibility of an origin of L3 in Asia was also proposed by Cabrera et al. (2018) based on the similar coalescence dates of L3 and its Eurasian-distributed M and N derivative clades (ca. 70 kya), the distant location in

Southeast Asia Southeast Asia, also spelled South East Asia and South-East Asia, and also known as Southeastern Asia, South-eastern Asia or SEA, is the geographical south-eastern region of Asia, consisting of the regions that are situated south of mainland ...

of the oldest known subclades of M and N, and the comparable age of the paternal haplogroup DE. According to this hypothesis, after an initial out-of-Africa migration of bearers of pre-L3 (L3'4*) around 125 kya, there would have been a back-migration of females carrying L3 from Eurasia to East Africa sometime after 70 kya. The hypothesis suggests that this back-migration is aligned with bearers of paternal haplogroup E, which it also proposes to have originated in Eurasia. These new Eurasian lineages are then suggested to have largely replaced the old autochthonous male and female North-East African lineages. According to other research, though earlier migrations out of Africa of anatomically modern humans occurred, current Eurasian populations descend instead from a later migration from Africa dated between about 65,000 and 50,000 years ago (associated with the migration out of L3). Vai et al. (2019) suggest, from a newly discovered old and deeply-rooted branch of maternal haplogroup N found in early Neolithic North African remains, that haplogroup L3 originated in East Africa between 70,000 and 60,000 years ago, and both spread within Africa and left Africa as part of the Out-of-Africa migration, with haplogroup N diverging from it soon after (between 65,000 and 50,000 years ago) either in Arabia or possibly North Africa, and haplogroup M originating in the Middle East around the same time as N. A study by Lipson et al. (2019) analyzing remains from the Cameroonian site of

Shum Laka The archaeological site of Shum Laka is the most prominent rockshelter site in the Grasslands region of the Laka Valley, northwest Cameroon. Occupations at this rockshelter date to the Later Stone Age. This region is important to investigations of ...

found them to be more similar to modern-day

Pygmy peoples In anthropology, pygmy peoples are ethnic groups whose average height is unusually short. The term pygmyism is used to describe the phenotype of endemic short stature (as opposed to disproportionate dwarfism occurring in isolated cases in a ...

than to West Africans, and suggests that several other groups (including the ancestors of West Africans, East Africans and the ancestors of non-Africans) commonly derived from a human population originating in East Africa between about 80,000-60,000 years ago, which they suggest was also the source and origin zone of haplogroup L3 around 70,000 years ago.

Distribution

Interpolation maps for L3 total haplogroups

L3 is common in

Northeast Africa Northeast Africa, or ''Northeastern Africa'' or Northern East Africa as it was known in the past, is a geographic regional term used to refer to the countries of Africa situated in and around the Red Sea. The region is intermediate between North ...

and some other parts of East Africa, in contrast to others parts of Africa where the haplogroups L1 and L2 represent around two thirds of mtDNA lineages. L3 sublineages are also frequent in the

Arabian peninsula The Arabian Peninsula, (; ar, شِبْهُ الْجَزِيرَةِ الْعَرَبِيَّة, , "Arabian Peninsula" or , , "Island of the Arabs") or Arabia, is a peninsula of Western Asia, situated northeast of Africa on the Arabian Plat ...

. L3 is subdivided into several clades, two of which spawned the macrohaplogroups M and N that are today carried by most people outside Africa. There is at least one relatively deep non-M, non-N clade of L3 outside Africa, L3f1b6, which is found at a frequency of 1% in

Asturias Asturias (, ; ast, Asturies ), officially the Principality of Asturias ( es, Principado de Asturias; ast, Principáu d'Asturies; Galician-Asturian: ''Principao d'Asturias''), is an autonomous community in northwest Spain. It is coextensiv ...

, Spain. It diverged from African L3 lineages at least 10,000 years ago. According to Maca-Meyer et al. (2001), "L3 is more related to

Eurasia Eurasia (, ) is the largest continental area on Earth, comprising all of Europe and Asia. Primarily in the Northern and Eastern Hemispheres, it spans from the British Isles and the Iberian Peninsula in the west to the Japanese archipelag ...

n haplogroups than to the most divergent African clusters L1 and L2". L3 is the haplogroup from which all modern humans outside Africa derive. However, there is a greater diversity of major L3 branches within Africa than outside of it, the two major non-African branches being the L3 offshoots M and N.

Subclade distribution

Interpolation maps for L3b, L3d, L3e, L3f L3h, L3i, L3x and L3w haplogroups

L3 has seven equidistant descendants: L3a, L3b'f, L3c'd, L3e'i'k'x, L3h, M, N. Five are African, while two are associated with the Out of Africa event. * N –

possibly due to migration from Africa, and North Africa possibly due to back-migration from Eurasia. * M –

Asia Asia (, ) is one of the world's most notable geographical regions, which is either considered a continent in its own right or a subcontinent of Eurasia, which shares the continental landmass of Afro-Eurasia with Africa. Asia covers an are ...

, the

Mediterranean Basin In biogeography, the Mediterranean Basin (; also known as the Mediterranean Region or sometimes Mediterranea) is the region of lands around the Mediterranean Sea that have mostly a Mediterranean climate, with mild to cool, rainy winters and wa ...

, and parts of Africa due to back-migration. * L3a – East Africa. Moderate to high frequencies found among the Sanye, Samburu, Iraqw, Yaaku, El-Molo and other minor indigenous populations from the

East African Rift The East African Rift (EAR) or East African Rift System (EARS) is an active continental rift zone in East Africa. The EAR began developing around the onset of the Miocene, 22–25 million years ago. In the past it was considered to be part of ...

Valley. It is infrequent to nonexistent in Sudan and the Sahel zone. ** L3a1 – Found across Eastern Africa. Estimated age of 35.8–39.3 ka. ** L3a2 – Found across Eastern Africa. Estimated age of 48.3–57.7 ka.See Supplemental_TreeUpdatedOctober.xls under th
Supplementary data
of * L3b'f ** L3b – Spread from East Africa in the

upper paleolithic The Upper Paleolithic (or Upper Palaeolithic) is the third and last subdivision of the Paleolithic or Old Stone Age. Very broadly, it dates to between 50,000 and 12,000 years ago (the beginning of the Holocene), according to some theories coin ...

to West-Central Africa. Some subclades spread from Central Africa to East Africa with the Bantu migration. *** L3b1a – Common subclade. Estimated age of 11.7-14.8 ka. **** L3b1a2 – Subclade found in Northeast Africa, the Maghreb, and Middle East. Emerged 12–14 ka. ** L3f – Northeast Africa, Sahel, Arabian peninsula, Iberia. Gaalien, Beja *** L3f1 **** L3f1a – Carried by migrants from Eastern Africa into the Sahel and Central Africa. **** L3f1b – Carried by migrants from Eastern Africa into the Sahel and Central Africa. ***** L3f1b1 – Carried from Central Africa into Southern and Eastern Africa with the Bantu migration. ****** L3f1b1a – Settled from East-Central Africa to Central-West Africa and into North Africa and Berber regions. ***** L3f1b4 – Carried from Central Africa into Southern and Eastern Africa with the Bantu migration. **** L3f1b6 – Rare, found in Iberia. *** L3f2 – Primarily distributed in East Africa. Also found in North Africa and Central Africa. *** L3f3 – Spread from Eastern Africa to Chad and the Sahel around 8–9 ka. Found in the

Chad Basin The Chad Basin is the largest endorheic basin in Africa, centered on Lake Chad. It has no outlet to the sea and contains large areas of semi-arid desert and savanna. The drainage basin is roughly coterminous with the sedimentary basin of the sa ...

. * L3c'd ** L3c – Extremely rare lineage with only two samples found so far in Eastern Africa and the

Near East The ''Near East''; he, המזרח הקרוב; arc, ܕܢܚܐ ܩܪܒ; fa, خاور نزدیک, Xāvar-e nazdik; tr, Yakın Doğu is a geographical term which roughly encompasses a transcontinental region in Western Asia, that was once the hist ...

. ** L3d – Spread from East Africa in the

to Central Africa. Some subclades spread to East Africa with the Bantu migration. Found among the

Fulani The Fula, Fulani, or Fulɓe people ( ff, Fulɓe, ; french: Peul, links=no; ha, Fulani or Hilani; pt, Fula, links=no; wo, Pël; bm, Fulaw) are one of the largest ethnic groups in the Sahel and West Africa, widely dispersed across the region. ...

Chad Chad (; ar, تشاد , ; french: Tchad, ), officially the Republic of Chad, '; ) is a landlocked country at the crossroads of North and Central Africa. It is bordered by Libya to the north, Sudan to the east, the Central African Republic ...

ians,

Ethiopian Ethiopians are the native inhabitants of Ethiopia, as well as the global diaspora of Ethiopia. Ethiopians constitute several component ethnic groups, many of which are closely related to ethnic groups in neighboring Eritrea and other parts of ...

Akan people The Akan () people live primarily in present-day Ghana and Ivory Coast in West Africa. The Akan language (also known as ''Twi/Fante'') are a group of dialects within the Central Tano branch of the Potou–Tano subfamily of the Niger–Con ...

Mozambique Mozambique (), officially the Republic of Mozambique ( pt, Moçambique or , ; ny, Mozambiki; sw, Msumbiji; ts, Muzambhiki), is a country located in southeastern Africa bordered by the Indian Ocean to the east, Tanzania to the north, Malawi ...

Yemen Yemen (; ar, ٱلْيَمَن, al-Yaman), officially the Republic of Yemen,, ) is a country in Western Asia. It is situated on the southern end of the Arabian Peninsula, and borders Saudi Arabia to the north and Oman to the northeast and ...

ites,

Egyptians Egyptians ( arz, المَصرِيُون, translit=al-Maṣriyyūn, ; arz, المَصرِيِين, translit=al-Maṣriyyīn, ; cop, ⲣⲉⲙⲛ̀ⲭⲏⲙⲓ, remenkhēmi) are an ethnic group native to the Nile, Nile Valley in Egypt. Egyptian ...

Berbers , image = File:Berber_flag.svg , caption = The Berber ethnic flag , population = 36 million , region1 = Morocco , pop1 = 14 million to 18 million , region2 = Algeria , pop2 ...

*** L3d3a1 – Primarily found in Southern Africa. Supplementary data . * L3e'i'k'x ** L3e – Spread from East Africa in the

to West-Central Africa. It is the most common L3 sub-clade in

Bantu Bantu may refer to: *Bantu languages, constitute the largest sub-branch of the Niger–Congo languages *Bantu peoples, over 400 peoples of Africa speaking a Bantu language * Bantu knots, a type of African hairstyle * Black Association for Nationa ...

-speaking populations. L3e is suggested to be associated with a Central African origin and is also the most common L3 subclade amongst African Americans,

Afro-Brazilian Afro-Brazilians ( pt, afro-brasileiros; ) are Brazilians who have predominantly African ancestry (see " preto"). Most members of another group of people, multiracial Brazilians or ''pardos'', may also have a range of degree of African ancestry. ...

s and

Caribbean The Caribbean (, ) ( es, El Caribe; french: la Caraïbe; ht, Karayib; nl, De Caraïben) is a region of the Americas that consists of the Caribbean Sea, its islands (some surrounded by the Caribbean Sea and some bordering both the Caribbean ...

s *** L3e1 – Spread from West-Central Africa to Southwest Africa with the Bantu migration. Found in

Angola , national_anthem = "Angola Avante"() , image_map = , map_caption = , capital = Luanda , religion = , religion_year = 2020 , religion_ref = , coordinat ...

(6.8%). Mozambique,

Sudan Sudan ( or ; ar, السودان, as-Sūdān, officially the Republic of the Sudan ( ar, جمهورية السودان, link=no, Jumhūriyyat as-Sūdān), is a country in Northeast Africa. It shares borders with the Central African Republic t ...

ese and

Kikuyu Kikuyu or Gikuyu (Gĩkũyũ) mostly refers to an ethnic group in Kenya or its associated language. It may also refer to: *Kikuyu people, a majority ethnic group in Kenya *Kikuyu language, the language of Kikuyu people *Kikuyu, Kenya, a town in Centr ...

Kenya ) , national_anthem = " Ee Mungu Nguvu Yetu"() , image_map = , map_caption = , image_map2 = , capital = Nairobi , coordinates = , largest_city = Nairobi , ...

, as well as in

, the

Tikar The Tikar (also Tikari, Tige, Tigar, Tigre, Tikali) are a central African people who inhabit the Western High Plateau in Cameroon. They are known as great artisans and storytellers. Once a nomadic people, some oral traditions trace the origin of ...

Cameroon Cameroon (; french: Cameroun, ff, Kamerun), officially the Republic of Cameroon (french: République du Cameroun, links=no), is a country in west-central Africa. It is bordered by Nigeria to the west and north; Chad to the northeast; the ...

, and among the

of Ghana. *** L3e5 – Originated in the Chad Basin. Found in

Algeria ) , image_map = Algeria (centered orthographic projection).svg , map_caption = , image_map2 = , capital = Algiers , coordinates = , largest_city = capital , relig ...

,
S5 Table
/ref> as well as

Burkina Faso Burkina Faso (, ; , ff, 𞤄𞤵𞤪𞤳𞤭𞤲𞤢 𞤊𞤢𞤧𞤮, italic=no) is a landlocked country in West Africa with an area of , bordered by Mali to the northwest, Niger to the northeast, Benin to the southeast, Togo and Ghana t ...

Nigeria Nigeria ( ), , ig, Naìjíríyà, yo, Nàìjíríà, pcm, Naijá , ff, Naajeeriya, kcg, Naijeriya officially the Federal Republic of Nigeria, is a country in West Africa. It is situated between the Sahel to the north and the Gulf o ...

, South

Tunisia ) , image_map = Tunisia location (orthographic projection).svg , map_caption = Location of Tunisia in northern Africa , image_map2 = , capital = Tunis , largest_city = capital , ...

, South

Morocco Morocco (),, ) officially the Kingdom of Morocco, is the westernmost country in the Maghreb region of North Africa. It overlooks the Mediterranean Sea to the north and the Atlantic Ocean to the west, and has land borders with Algeria to A ...

and

Egypt Egypt ( ar, مصر , ), officially the Arab Republic of Egypt, is a List of transcontinental countries, transcontinental country spanning the North Africa, northeast corner of Africa and Western Asia, southwest corner of Asia via a land bridg ...

** L3i Almost exclusively found in East Africa. *** L3i1 **** L3i1b – Subclade is found in Yemen, Ethiopia, and among Gujarati Indians. *** L3i2 (former L3w) – Found in the Horn of Africa and Oman. ** L3k – Rare haplogroup primarily found in North Africa and the Sahel. ** L3x – Almost exclusively found in East Africa. Found among Ethiopian

Oromos The Oromo (pron. Oromo: ''Oromoo'') are a Cushitic ethnic group native to the Oromia region of Ethiopia and parts of Northern Kenya, who speak the Oromo language (also called ''Afaan Oromoo'' or ''Oromiffa''), which is part of the Cushitic b ...

GUR46 on table 1. is a mtDNA haplogroup L3x2a. * L3h – Almost exclusively found in East Africa. ** L3h1 – Primarily found in East Africa with branches of L3h1b1 sporadically found in the Sahel and North Africa. ** L3h2 – Found in Northeast Africa and Socotra. Split from other L3h branches as early as 65–69 ka during the

middle paleolithic The Middle Paleolithic (or Middle Palaeolithic) is the second subdivision of the Paleolithic or Old Stone Age as it is understood in Europe, Africa and Asia. The term Middle Stone Age is used as an equivalent or a synonym for the Middle Paleol ...

Ancient and historic samples

Haplogroup L3 has been observed in an ancient fossil belonging to the

Pre-Pottery Neolithic B Pre-Pottery Neolithic B (PPNB) is part of the Pre-Pottery Neolithic, a Neolithic culture centered in upper Mesopotamia and the Levant, dating to years ago, that is, 8800–6500 BC. It was typed by British archaeologist Kathleen Kenyon durin ...

culture. L3x2a was observed in a 4,500 year old hunter-gather excavated in Mota, Ethiopia, with the ancient fossil found to be most closely related to modern Southwest Ethiopian populations. Haplogroup L3 has also been found among ancient Egyptian mummies (1/90; 1%) excavated at the

Abusir Abusir ( ar, ابو صير ; Egyptian ''pr wsjr'' cop, ⲃⲟⲩⲥⲓⲣⲓ ' "the House or Temple of Osiris"; grc, Βούσιρις) is the name given to an Egyptian archaeological locality – specifically, an extensive necropolis ...

el-Meleq archaeological site in Middle Egypt, with the rest deriving from Eurasian subclades, which date from the Pre- Ptolemaic/late

New Kingdom New is an adjective referring to something recently made, discovered, or created. New or NEW may refer to: Music * New, singer of K-pop group The Boyz Albums and EPs * ''New'' (album), by Paul McCartney, 2013 * ''New'' (EP), by Regurgitator ...

and Ptolemaic periods. The Ancient Egyptian mummies bore Near eastern genomic component most closely related to modern near easterners. Additionally, haplogroup L3 has been observed in ancient Guanche fossils excavated in

Gran Canaria Gran Canaria (, ; ), also Grand Canary Island, is the third-largest and second-most-populous island of the Canary Islands, an archipelago off the Atlantic coast of Northwest Africa which is part of Spain. the island had a population of that ...

and

Tenerife Tenerife (; ; formerly spelled ''Teneriffe'') is the largest and most populous island of the Canary Islands. It is home to 43% of the total population of the Archipelago, archipelago. With a land area of and a population of 978,100 inhabitant ...

on the

Canary Islands The Canary Islands (; es, :es:Canarias, Canarias, ), also known informally as the Canaries, are a Spanish Autonomous communities of Spain, autonomous community and archipelago in the Atlantic Ocean, in Macaronesia. At their closest point to ...

, which have been radiocarbon-dated to between the 7th and 11th centuries CE. All of the clade-bearing individuals were inhumed at the Gran Canaria site, with most of these specimens found to belong to the L3b1a subclade (3/4; 75%) with the rest from both islands (8/11; 72%) deriving from Eurasian subclades. The Guanche skeletons also bore an autochthonous Maghrebi genomic component that peaks among modern

Berbers , image = File:Berber_flag.svg , caption = The Berber ethnic flag , population = 36 million , region1 = Morocco , pop1 = 14 million to 18 million , region2 = Algeria , pop2 ...

, which suggests that they originated from ancestral Berber populations inhabiting northwestern Affoundnat a high ncy A variety of L3 have been uncovered in ancient remains associated with the Pastoral Neolithic and Pastoral Iron Age of East Africa.

Tree

This

phylogenetic tree A phylogenetic tree (also phylogeny or evolutionary tree Felsenstein J. (2004). ''Inferring Phylogenies'' Sinauer Associates: Sunderland, MA.) is a branching diagram or a tree showing the evolutionary relationships among various biological spec ...

of haplogroup L3 subclades is based on the paper by Mannis van Oven and Manfred Kayser ''Updated comprehensive phylogenetic tree of global human mitochondrial DNA variation'' and subsequent published research. Most Recent Common Ancestor (MRCA) * L1-6 ** L2-6 *** L2'3'4'6 **** L3'4'6 ***** L3'4 ****** L3 ******* L3a ******** L3a1 ********* L3a1a ********* L3a1b ******** L3a2 ********* L3a2a ******* L3b'f ******** L3b ********* L3b1 ********** L3b1a *********** L3b1a1 *********** L3b1a2 *********** L3b1a3 *********** L3b1a4 *********** L3b1a5 ************ L3b1a5a ************ L3b1a6 ************ L3b1a7 ************* L3b1a7 ************ L3b1a8 ************ L3b1a9 ************* L3b1a9a *********** L3b1a10 *********** L3b1a11 ********** L3b1b *********** L3b1b1 ********* L3b2 ********** L3b2a ********** L3b2a ********* L3b3 ******** L3f ********* L3f1 ********** L3f1a *********** L3f1a1 ********** L3f1b *********** L3f1b1 *********** L3f1b2 ************ L3f1b2a ************ L3f1b3 ************ L3f1b4 ************* L3f1b4a ************** L3f1b4a1 ************* L3f1b4b ************* L3f1b4c *********** L3f1b5 ********* L3f2 ********** L3f2a ********** L3f2b ********* L3f3 ********** L3f3a ********** L3f3b ******* L3c'd ******** L3c ******** L3d ********* L3d1-5 ********** L3d1 *********** L3d1a ************ L3d1a1 ************* L3d1a1a *********** L3d1b ************ L3d1b1 *********** L3d1c *********** L3d1d ********** 199 *********** L3d2 *********** L3d5 ********** L3d3 *********** L3d3a ********** L3d4 ********** L3d5 ******* L3e'i'k'x ******** L3e ********* L3e1 ********** L3e1a *********** L3e1a1 ************ L3e1a1a *********** 152 ************ L3e1a2 ************ L3e1a3 ********** L3e1b ********** L3e1c ********** L3e1d ********** L3e1e ********* L3e2 ********** L3e2a *********** L3e2a1 ************ L3e2a1a ************ L3e2a1b ************* L3e2a1b1 ********** L3e2b *********** L3e2b1 ************ L3e2b1a *********** L3e2b2 *********** L3e2b3 ********* L3e3'4'5 ********** L3e3'4 *********** L3e3 ************ L3e3a ************ L3e3b ************* L3e3b1 ************ L3e4 ********** L3e5 ******** L3i ********* L3i1 ********** L3i1a ********** L3i1b ********* L3i2 ******** L3k ********* L3k1 ******** L3x ********* L3x1 ********** L3x1a *********** L3x1a1 *********** L3x1a2 ********** L3x1b ********* L3x2 ********** L3x2a *********** L3x2a1 ************ L3x2a1a ********** L3x2b ******* L3h ******** L3h1 ********* L3h1a ********** L3h1a1 ********** L3h1a2 *********** L3h1a2a *********** L3h1a2b ********* L3h1b ********** L3h1b1 *********** L3h1b1a ************ L3h1b1a1 ********** L3h1b2 ******** L3h2 ******* M ******* N

References

Notes

{{Reflist, group=Note

External links

* General ** Ian Logan'
Mitochondrial DNA Site
* Haplogroup L3 ** Mannis van Oven'

*

from ''

National Geographic ''National Geographic'' (formerly the ''National Geographic Magazine'', sometimes branded as NAT GEO) is a popular American monthly magazine published by National Geographic Partners. Known for its photojournalism, it is one of the most widel ...

'' L3 Recent African origin of modern humans