Haplogroup L3 (mtDNA)
Haplogroup L3 is a human mitochondrial DNA haplogroup. The clade has played a pivotal role in the early dispersal of anatomically modern humans.
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 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. 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 60–35,000 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 roughly 110 kya, and the L3'4 clade at 95 kya.
The possibility of an origin of L3 already in Asia was also proposed by Cabrera et al. based on the similar coalescence dates of L3 and its Eurasian-distributed M and N derivative clades, the distant location in Southeast Asia of the oldest 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 around 125 kya, there would have been a back-migration of females carrying L3 from Eurasia to East Africa sometime after 70 kya. It is suggested that this back-migration is aligned with bearers of paternal haplogroup E, which is also proposed 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. Vai et al. 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 70,000-60,000 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 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 analyzing remains from the Cameroonian site of Shum Laka found them to be more similar to modern-day Pygmy peoples than to West Africans, and suggests that several other groups 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
L3 is common in Northeast Africa 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.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, Spain. It diverged from African L3 lineages at least 10,000 years ago.
According to Maca-Meyer et al., "L3 is more related to Eurasian 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
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 – Eurasia and parts of Africa due to back-migration.
- M – Asia, the Mediterranean Basin, 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 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.
- L3b'f
- * L3b – Spread from East Africa in the upper paleolithic 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.
- L3c'd
- * L3c – Extremely rare lineage with only two samples found so far in Eastern Africa and the Near East.
- * L3d – Spread from East Africa in the upper paleolithic to Central Africa. Some subclades spread to East Africa with the Bantu migration. Found among the Fulani, Chadians, Ethiopians, Akan people, Mozambique, Yemenites, Egyptians, Berbers
- ** L3d3a1 – Primarily found in Southern Africa.
- L3e'i'k'x
- * L3e – Spread from East Africa in the upper paleolithic to West-Central Africa. It is the most common L3 sub-clade in Bantu-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-Brazilians and Caribbeans
- ** L3e1 – Spread from West-Central Africa to Southwest Africa with the Bantu migration. Found in Angola. Mozambique, Sudanese and Kikuyu from Kenya as well as in Yemen and among the Akan people
- ** L3e5 – Originated in the Chad Basin. Found in Algeria, as well as Burkina Faso, Nigeria, South Tunisia, South Morocco and Egypt
- * L3i Almost exclusively found in East Africa.
- ** L3i1
- *** L3i1b – Subclade is found in Yemen, Ethiopia, and among Gujarati Indians.
- ** L3i2 – 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, Egyptians
- 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.
Ancient and historic samples
A variety of L3 have been uncovered in ancient remains associated with the Pastoral Neolithic and Pastoral Iron Age of East Africa.
Culture | Genetic cluster or affinity | Country | Site | Date | Maternal Haplogroup | Paternal Haplogroup | Source |
Early pastoral | PN | Kenya | Prettejohn's Gully | 4060–3860 | L3f1b | – | Prendergast 2019 |
Pastoral Neolithic | PN | Kenya | Cole’s Burial | 3350–3180 | L3i2 | E-V32 | Prendergast 2019 |
Pastoral Neolithic or Elmenteitan | PN | Kenya | Rigo Cave | 2710–2380 | L3f | E-M293 | Prendergast 2019 |
Pastoral Neolithic | PN | Kenya | Naishi Rockshelter | 2750–2500 | L3x1a | E-V1515 | Prendergast 2019 |
Pastoral Neolithic | PN | Tanzania | Gishimangeda Cave | 2490–2350 | L3x1 | – | Prendergast 2019 |
Pastoral Neolithic | PN | Kenya | Naivasha Burial Site | 2350–2210 | L3h1a1 | E-M293 | Prendergast 2019 |
Pastoral Neolithic | PN | Kenya | Naivasha Burial Site | 2320–2150 | L3x1a | E-M293 | Prendergast 2019 |
Pastoral Neolithic | PN | Tanzania | Gishimangeda Cave | 2150–2020 | L3i2 | E-M293 | Prendergast 2019 |
Pastoral Neolithic or Elmenteitan | PN | Kenya | Njoro River Cave II | 2110–1930 | L3h1a2a1 | – | Prendergast 2019 |
Pastoral Neolithic | N/A | Tanzania | Gishimangeda Cave | 2000–1900 | L3h1a2a1 | – | Prendergast 2019 |
Pastoral Neolithic | PN | Kenya | Ol Kalou | 1810–1620 | L3d1d | E-M293 | Prendergast 2019 |
Pastoral Iron Age | PIA | Kenya | Kisima Farm, C4 | 1060–940 | L3h1a1 | E-M75 | Prendergast 2019 |
Pastoral Iron Age | PIA | Kenya | Emurua Ole Polos | 420–160 | L3h1a1 | E-M293 | Prendergast 2019 |
Pastoral Iron Age | PN outlier | Kenya | Kokurmatakore | N/A | L3a2a | E-M35 | Prendergast 2019 |
Tree
This phylogenetic tree 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
- 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