Monday, June 11, 2012
Haplogroup L3(M,N) Expanded before the Out of Africa Exit
L3(M,N) probably spread across Africa before the Out of Africa event that led to the dispersal of anatomically modern humans to Eurasia. Sores et al (2012) admit that the several branches of L3 probably expanded soon after the emergence of L3, they fail to comprehend the full extent of this finding. If we are using the Toba super-eruptiobn of 73.5ka as a baseline blocking any OoA event of amh (Oppenheimer,2012; Sores et al, 2012), L(M,N) must have spread across Africa before the OoA event.
Sores et al (2012) believes that mtDNA haplogroups M and N originated in Eurasia. Sores et al (2012) founded this conclusion on Olivieri et al (2006) who maintained that around 50kya, M1 entered Africa as a result of a bacxk migration from Eurasia; and a sample limited to Sudanese, Ethiopians and Somalis.
Sores et al (2012) argues that L3(M,N and R) originated in Eurasia, based on Olivieri et al (2006). Olivieri et al (2006) suggest that haplogroup M1 probably did not originate in Africa before the out of Africa exit . They opine that M1 probably represents a back-migration into Ethiopia . Olivieri et al base this conclusion on :
1) the absence of any distinguishing M1 root mutations in Asian M haplogroups ;
2) the presence of M1 only in East Africa and North Africa; and
3) the lack of any Asian specific clades within M1
Olivieri et al (2006) argue that M1 was probably spread from the Levant back into Africa by the Aurignacian culture. The craniofacial and molecular evidence does not support this conclusion (Winters, 2007,2008b).
Gonzalez et al (2007) and Olivieri et al (2006) have assumed that haplogroup L3(M,N), was probably carried to western Eurasia via the Levant. But the archaeological and craniometric eveidence indicates that the Levant was still occupied by Neanderthal man until 32kya. The archaeological evidence indicates that Around 40kya Europe was still occupied mainly by Neanderthals.
The archaeological record informs us that Cro-Magnon people belonging to the Aurignacian culture carried hg N (Caramelli et al , 2003). They replaced the Neanderthal population of the Levant, at Ksar Akil around 32, 000 years ago (Gilead, 2005; Steven, 2001). This was 10k after Cro-Magnon people had settled Iberia.
Anatomically modern humans replaced in Europe around 32,000 by the CroMagnon people at Les Eyzies in France. It is also evident that archaic humans were replaced in much of the Levant by the Levantine Aurignacian culture bearers by a local variant of this technology at Ksar Akil Xlll-Vll 32kya , not 60-50kya. The archaeological evidence makes a back migration from the Levant.
Oppenheimer (2012) makes it clear that amh colonization of the Levant failed; and by60kya Neanderthal populations dominated the Levant.
Clearly, the dates for L3(M,N) in western Eurasian are incongruent to TMRCA of the populations carrying the L3(M,N) lineages into eastern Eurasia which probably date to 60-65kya. This incongruence in relation to the dates for this haplogroup in eastern Eurasia, and its complete absence in much of western Eurasia today suggest that the population carrying this gene into Eurasia may not have entered Eurasian during the recognized Africa exit event.
There is considerable evidence that M1 is found in Asia. Researchers have found the M1 haplogroup in the Caucasus ( Bermisheva et al, 2004; Tambets et al, 2000), Central Asia, and East Asia (Comas et al, 1998; Fucharoen et al, 2001). In addition, the Russian haplotype 16183c-16189 ,16249, 16311 match the M1 HVSI sequence (Malyarchuk et al, 2004).
Even though Olivieri et al (2007) claim that East African M1 root mutations are absent in Eurasian M sister clades is not supported by the evidence. For example researchers have found that the Tanzanian M1 haplogroup cluster with people from Oceania (Gonder et al, 2006). And, as mentioned earlier the M1 mutations 16129,16189,16249 and 16311 are found in many southeast and East Asian haplogroups (Fucharoen et al, 2001; Yao et al, 2002).
It is also not true that HG M1 is absent in India. Kivisild et al noted that 26 of the subjects in their study belonged to the M1 haplogroup.(7) These researchers reported sub-cluster M1 was found mainly in Kerala and Karnataka high caste individuals.(7)
It is clear that the molecular evidence does not support Olivieri et al (2006) hypothesis that M1 is probably the result of a back migration. This evidence on the other hand confirms the hypothesis of Quintana-Murci et al (1999) that M1 was probably already present in East Africa when the out of Africa exit/ event took place.
The molecular evidence makes it clear that haplogroup M1 is not confined solely to Ethiopia as maintained by Olivieri et al (2006). This haplogroup along with HGs N and M*, are also found in Tanzania, Uganda, Egypt and the Senegambian region (Gonzalez et al, 2006; Gonder et al, 2006; Winters, 2007). In Tanzania the predominate M1 clades are M1 , M1a1 and M1a5. In Senegal the predominate M1 lineage is M1c1.
In addition to M1 in Africa, we also find haplogroups M*, M23, M3 positions 482 and 16126; M30 positions 195A and 15431; and M33 position 2361. It is interesting to note that the presence of these genes, which are normally found in India are also found in Africa, is interesting given the presence of M1 in India and the existence of these genes among populations stretching from Africa into Yemen on into India along a path associated with the spread of the Tihama culture (Winters, 2008) .
In addition to haplogroups M1, M* and N in Sub-Saharan Africa we also find among the Senegambians hapotype AF24 (DQ112852) , which is delineated by a DdeI site at 10394 and AluI site of np 10397. The AF-24 haplotype is a branch of the African subhaplogroup L3 (Chen, 2000). This is the same delineation of haplogroup M*. It is clear from the molecular evidence that the M1, M and N haplogroups are found not only in Northeast Africa, but across Africa from East to West (Winters, 2007).
Haplogroup LOd is found at the root of human mtDNA. Gonder et al (2006) maintains that LOd is “the most basal branch of the gene tree”. The TMRCA for LOd is 106kya. This makes haplotype AF-24 much older than L3a and probably explains why this haplotype is found among the Khwe/Khoisan (Chen et al,2000).
The TMRCA of LOd dates to 106kya. As a result, anatomically modern humans (amh) had plenty of time to spread this haplogroup to Senegal. In West Africa the presence of amh date to the Upper Palaeolithic (Giresse,2008). The archaeological evidence makes it clear that amh had ample opportunity to spread LOd and L3(M,N) which has an affinity to AF-24 (Chen,2000), to West Africa during this early period of demic diffusion of amh in Africa.
The earliest evidence of human activity in West Africa is typified by the Sangoan industry (Phillipson,2005). The amh associated with the Sangoan culture may have deposited Hg LOd and haplotype AF-24 in Senegal thousands of years before the exit of amh from Africa. This is because it was not until 65kya that the TMRCA of non-African L3(M,N) exited Africa (Kivisild et al, 2006).
Anatomically modern humans arrived in Senegal during the Sangoan period. Sangoan artifacts spread from East Africa to West Africa between 100-80kya. In Senegal Sangoan material has been found near Cap Manuel (Giresse, 2008), Gambia River in Senegal (Davies,1967; Wai-Ogussu,1973); and Cap Vert (Phillipson,2005).
In conclusion, because the Neanderthal dominated the Levant when the imagined back migration of M1 occurred 50kya ,we must reject the contention of Gonzalez et al. (2007) and Olivieri et al. (2007) and Sores et al (2012) that M1 originated in Asia because 1) the possible Senegalese origin of the M1c subclade; 2) the absence of the AF-24 haplotype of haplogroup LOd in Asia; and 3) the African origin of the Dravidian speakers of India (2007,2008)who carry the most diverse M haplogroups.
Moreover, the existence of the L3a(M) motif in the Senegambia characterized by the DdeI site np 10394 and AluI site np 10397 in haplotype AF24 (DQ112852) make a ‘back migration of M1 to Africa highly unlikely, because of the ancientness of this haplotype. The first amh to reach Senegal belonged to the Sangoan culture which spread from East Africa to West Africa probably between 100-80kya.
The presence of the AF-24 is a haplotype of haplogroup LOd makes it clear that this haplotype is not only an ancient human genome. It is also evidence that AF-24 probably did not originate in Asia, since AF-24 was found among the Senegalese and Khoisan.
This reflects an early migration from East Africa to West Africa. The presence of basal nucleotides characteristic of macrohaplogroup L3(M) in West Africa and the reality that M1 does not descend from an Asian M macrohaplogroup because of the absence of AF24 in Asia (Sun et al, 2005) and its presence among the Khoisan and Senegalese suggest that expansion of M1 was probably from Africa to Eurasia. The existence of haplotype AF-24 and basal L3(M) lineage in East and West Africa suggest the probable existence of the Proto-M1 lineage in Africa, not Eurasia before haplogroup L3(M,N) carriers exited Africa.
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