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Thursday, March 11, 2010

The early expansion of H. sapiens in Africa (mtDNA)


As you probably know, I have been working as of late on the puzzle of the early human mtDNA, in the context of Africa, using particularly
Behar 2008 as reference.

I published some stuff previously (L0, L1, L2 & L5, L3'4'6 and considerations on Arabian L(xM,N)) and went into a lengthy discussion with Terry about where exactly did the various L lineages coalesce and, of course, human prehistoric boating abilities (an impossible for him before the arrival to almost mythical archipelago of Wallacea, aka Eastern Indonesia).

So I took my time to estimate the composite centroids of each lineage, at every level, from bottom up. Some centroids are plainly valid as likely origins for the major clades, while a few others are more questionable (see below). After a subjective but common-sensical revision of these I came up with the following maps that reconstruct early human expansion:


First expansion:

There seem to be basically two moments of demographic expansion, the first one happened at about 9 (coding region) mutations downstream of the most recent common ancestor (mitochondrial Eve), where the nodes of L0, L1 and L2"6 are in the phylogenetic tree, maybe corresponding with some favorable conditions, which are hard to unravel.


This map shows the earliest expansion at the very phylogenetic origin of the species (purple arrow) and the subsequent three populations generated, defined by the mtDNA lineages L0, L1 and L2"6, all three showing some sign of expansion around the 9th CR mutation (i.e. approximately at the same time, if the molecular clock makes any sense whatsoever).

Of course, L1 and L2"6 shared phylogeny for 5 CR mutations (L1"6), what seems to mean that they had reached Central Africa by that time.

The location that may cause more controversy is that of L0 around Lake Tanganyika, because this lineage is most commonly associated with Khoisan people. However only the L0d1'2 subslineage is totally exclusive of this macro-ethnicity and other L0 lineages are scattered towards the Nile and even as far as Kuwait. The overall centroid, as well as those of L0d and L0a'b'f'k, showed up around there, so I decided that it stands that way, even if admittedly origins around Ethiopia-Sudan or Southern Africa can also be argued for (as did Doron Behar).


Second expansion:

The second major expansion is located at around the 20th CR mutation downstream from "Eve" and in my opinion may be correlated with the Abbassia Pluvial, some 120-90,000 years ago, which probably improved living conditions allowing for significative demographic growth.

It is also the prelude of the migration out of Africa into Asia.


At this moment we see L0 finally splitting up clearly between a northern and a southern group, the latter leading to the Khoisan peoples.

We also see signal of expansion of L1c (but not yet L1b), which correlates best with Pygmies (and also some other peoples of the jungle belt essentially).

And we see the split of L2'3'4'6 (the branching of L5 is earlier, just marked it for the record), first between Western (L2) and Eastern (L3'4'6) populations, centered at Central Africa and what seems to be Eritrea respectively. Then L3'4 split up, with L3 expanding rapidly in Ethiopia, Sudan and towards Lake Chad and L4 scattering along the Rift Valley down to Tanzania, where it is still the most important lineage among the Hadza and Sandawe. L3, of course, also expanded eastward towards Asia where its sublineages M and N would find enormous opportunities.

The minor lineage L6 would not expand till a later moment, so at this time it was yet some "private" L3'4'6*. I suspect that it also benefited from the Out of Africa migration, along with L4b and some L0 subclades, because it's highest basal diversity is in Yemen, suggesting it coalesced there.

A word of caution must be said about the area of origin of L2. The actual composite centroid happens to be at the Niger but this location is the product of basal sublineage L2e, which is only represented by one individual in Behar's data, who happens to live at Guinea Bissau. But the major subclade, only diverged from L2 by a single CR mutation (unlike L2e, which has a very long stem), is L2a"d and this one has a clear center at Chad and, secondarily, the CAR.

Similarly, I ended up with a composite centroid at Yemen for L3 after considering M and N. But I can't ignore the absence of basal L3 sublineages in South Arabia, so I decided that the result without M and N is much more likely to reflect the reality.


The actual centroids:

Prior to the above maps, and after hard work, I came up with this preliminary map. The composite centroids here are the "raw" ones, without further consideration:



But, as said above, I have serious caveats about L2, L3 and L6 specially. I think that the areas depicted in the other maps reflect better the likely reality, with L2 coalescing at Chad, L3 at Sudan and L6 most probably at Yemen after the out-of-Africa episode.

As said before, I also have some caveats about the coalescence area of L0 but I can't come up with a clear alternative (it'd be either Ethiopia or Southern Africa but both weight about the same), so I left it that way.

Of course, the ultimate place of origin of Humankind (Homo sapiens), or at least of the most recent common ancestor (mtDNA Eve), remains a mystery. But guess that somewhere in the vast geography of Eastern Africa is a quite reasonable conjecture (Southern and Central Africa are also possibilities).

_______________

Erratum: there is an error with minor lineage L5 (thanks to Terry for noticing) but luckily doesn't affect much the overall picture. I made a meaningful error when calculating the centroid of L5a, what, in the maps here pushes the centroid of L5 significantly but not dramatically to the south, to South Sudan at the border with SW Ethiopia to be precise. In turn this affects the centroid of L2"6, L1"6 and "Eve", pushing them also proportionately southwards (not too much but something anyhow). The rest of the nodes are not affected, as they depend only of the downstream geography.

I think it's no big deal but I might correct the maps later on... if my perfectionist side prevails over my lazy one.

33 comments:

Maju said...

Replying to Terry's misplaced questions (see this discussion).

"But the south and north of what? My guess is somewhere in the grassy savannah in the southern Congo Basin".

Doesn't work. Uganda, or rather the border area between Uganda, Sudan, Congo and CAR fits best.

If L0 coalesced around Ethiopia, then Sudan fits best. Only if you make L0 coalesce in Southern Africa, then would your hypothesis make any sense. But considering that there's no apparent sign of expansion in that area, I seriously doubt it.

"Y-haps A and B have their centroids there, so we have a possible connection".

Those centroids don't have my quality seal of approval. I really don't make too much sense of them. But who knows?

Also it's Y-DNA...

"There seems next to be several expansions around the 9 mut. level. By then L1''6 had already diverged into L2''6 and L1. L1 had even diverged into L1b and L1c, and L5 had appeared."

L1"6 of course: already mentioned. L1 diverges precisely at the 9 mutations point and so does L1"6.

However L1b, L1c, L5 and L2"6 would only coalesce (branch out) after some more mutations. At this stage they are just L1* and L1"6*. L1c and L2"6 belong to the second map (around the 20th mutation). L5 is intermediate but being the only one in that parenthesis (and a small lineage) I did not consider useful to create a map only for it.

"At 9 mut in the south L0d managed to move further south from the Congo Basin, possibly into the drier savannah. L0a,b,f,k started its move north, possibly through a similar habitat".

In my draft centroid map, L0d and L0a'b'f'k are exactly south and north of the Tanganyika L0 centroid: at Malawi and the Serengeti respectively.

However it could be argued that I should have discarded that L0d3 from Kuwait but, as you can see in the results everything goes smoother without such discard, that admittedly "delays" the arrival to Southern Africa to the L0d1'2 stage.

But it makes good sense anyhow, because the southern L0 group has some La'b'f'k lineages and the northern L0 group has some L0d ones (at least in Kuwait).

The alternative is to discard Kuwait and Yemen (and in general all Arabian lineages) but at this moment I feel it's not correct. It would push the origin of L0 to Southern Africa faster (as happens in the hypothesis Behar manages) but would then demand a separate migration of several lineages.

There are too many L0 lineages shared between Khoisan and East Africans (and Arabians) not to raise some eyebrows. This alternative seems to work well, only "allowing" real separation of the two populations at the moment when the lineages are clearly diverging into a south and north group.

I reverse-engineered the whole haplogroup so I may even be right. Who knows?

"L5 may have moved beyond the grassland savannah of the northeast Congo Basin".

L5 has a very clear centroid in Sudan, as do L5a (south) and L5b (north). I have no doubts whatsoever about L5 being essentially Sudanese.

Maju said...

"Where was L1'2'3'6?"

That haplogroup does not exist.

If you mean L1"6, including also L5, then between the L1 and L2"6 coalescence areas most likely. What's that? Central African Republic? Around the Ubangi river I'd say.

"In fact L5 had split at 13 mut into a northeastern haplogroup (beyond the Congo Basin, L5c), and a southwestern one (L5a) within the Congo Basin".

Took me a while to decipher this but I finally made sense of it. You are partly correct: it seems I forgot, in my hurry to finish the labyrinth, to account properly for the Mozambican L5a2, what pushes the centroid of L5a quite far to the South, near Rwanda. However the overall L5 node is only pushed a bit south, to Southern Sudan instead of Central Sudan, and therefore the L2"6 centroid is almost not moved (just a bit to the south too).

I'll comment on it on an "erratum note" note but (luckily) is not such a large error as to have to rework all the maps.

terryt said...

Great work Maju. I'd place L0 a little north of where you have in and L1''6 a litlle south. In other words I suspect they both came from within a relativel restricted region rather than being widespread from the word go.

"If L0 coalesced around Ethiopia, then Sudan fits best".

But that's assuming it coalesced in Ethiopia. If it didn't your argument doesn't hold.

"However L1b, L1c, L5 and L2"6 would only coalesce (branch out) after some more mutations. At this stage they are just L1* and L1"6*".

Call them that if you wish.

"the southern L0 group has some La'b'f'k lineages and the northern L0 group has some L0d ones (at least in Kuwait)".

The southern memebers of La'b'f'k are quite possibly lines that remained behind as the haplogroup moved north. They fit that scenario precisely. And the Kuwait L0d is very odd, and could have arrived at any time, especially as it's not differentiated from the South African haplogroup.

" It would push the origin of L0 to Southern Africa faster (as happens in the hypothesis Behar manages) but would then demand a separate migration of several lineages".

Not really. They peel off as the haplogroup moves north.

"L5 has a very clear centroid in Sudan"

And Sudan is precisely 'beyond the grassland savannah of the northeast Congo Basin'.

I believe the evidence fits a popualtion expanding through grassland savannah better than any coastal population. So I also sispect very strongly that the migration OoA also took place through similar habitat.

Maju said...

"I'd place L0 a little north of where you have in and L1''6 a litlle south. In other words I suspect they both came from within a relativel restricted region rather than being widespread from the word go".

It's possible. However, as I say in the article, L1"6 must have been in Central Africa (roughly) since mutation #5 or before, so I can't move that bloc much. L0 is highly mobile instead (from Ethiopia to Southern Africa) until the L0d1'2 and L0a'b phases.

Ethiopia as origin of L0 fits best with your reasoning of a "restricted region" for the "Eve" ancestral clan. This is basically option B in Behar's paper and, I understand, the one he favors most.

But I did not dare to go that far and found instead what seems a natural gradual disentangling process within L0 in East Africa.

I'd have to check the archaeology, but I think we don't have much MSA or human presence in Southern Africa before 100,000, do we?

Instead we have dates as old as 190 Ka in Ethiopia (Omo) and at 160 Ka in Ethiopia (Omo, Herto) and Morocco (Jebel Irhoud). One wonders if these last might be related to L2e.

"Call them that if you wish".

For the sake of discursive clarity and my own mental lucidity, a naming procedure is needed. And this implies that haplogroups don't exist until they do exist (until they branch out).

"The southern memebers of La'b'f'k are quite possibly lines that remained behind as the haplogroup moved north".

In my reconstruction, L0k migrates southwards, while L0d3 northwards. It all depends of what origin for L0 you prefer.

"And the Kuwait L0d is very odd, and could have arrived at any time, especially as it's not differentiated from the South African haplogroup".

It is a different lineage. There are cases in which the same lineage is found at two locations, there are cases where a haplogroup branching late shows up at two locations... but this is a haplogroup branching out early into a Kuwaiti and a Khoisan branch. I don't know where exactly it arrived to Kuwait but I don't have any reason to think that slave traders massively raided the Kalahari Desert. The pattern northwards fits what the apparent moves of other L0 lineages, so I'm content with the result.

Maju said...

"Not really. They peel off as the haplogroup moves north".

Khoi-Kuwait, San-Yemen, Oman/Ethiopia-SE Bantu, S. Africa-Algeria, S. Africa/Kenya-Morocco: how do you "peel off" that?

There are L0 clades that are northerner, there are those that are southerner and there are quite a few that are both. And this last bit is quite interesting, IMO, and what has lead me to this conclusion.

You can argue it all you wish: it's a matter of subjectivity and I don't think it will be solved without much more data and work - if it can be solved at all.

"And Sudan is precisely 'beyond the grassland savannah of the northeast Congo Basin'".

I was thinking you meant Katanga or Zambia, LOL. Guess it depends where do you look at Congo River from.

"I believe the evidence fits a popualtion expanding through grassland savannah better than any coastal population. So I also sispect very strongly that the migration OoA also took place through similar habitat".

Well, the L3'4'6 group seems to have been specialized in coastal exploitation (and remember the remains we do have from Eritrea precisely) and I suspect it was also the case with some L0 groups, L0f in particular (centroid at Dar-es-Salaam) and, considering Blombos and related South African sites, other L0 groups such L0k and or L0d1'2 probably too.

L0 is one of the lineages that, curiously, appears more commonly at Arabia (L0k2, L0f2 and L0a1c have their centroids there). And remember that the Jawalpuram toolkit is very close to South African MSA and not really to East African MSA.

This could fit with whatever scenario for the origin of L0 but suggests that they were at least partly specialized in coastal (and lake?) activities and that they participated in the OoA, even if none of their lineages made it to India or survived the Toba scenario.

I don't reject savanna expansions either but both are highly compatible in my mind, specially as soon they would become several groups. The groups with coastal/lake best exploitation abilities would also have good chances of survival and expansion, as these are extremely productive environments.

terryt said...

"I was thinking you meant Katanga or Zambia, LOL".

They're northeast of the Congo Basin? Maju you really do need a new map.

"There are L0 clades that are northerner, there are those that are southerner and there are quite a few that are both".

Check out your own diagram. The first split is L0a'b'f'k and L0d. L0d is virtually confined to South Africa. The presence of L0d3 in Kuwait is hardly evidence for an ancient presence there.

The next split is L0a'b'f and L0k. L0k is also South African with a mysterious clade in Yemen at 33 mutations, so we can ignore its presence there as irelevant to L0's early expansion.

The next split is L0a'b and L0f. L0f is a little further north, found in South Africa and Ethiopia (Dar-es-Salaam you say). Again with a mysterious Oman clade.

The next split is L0a and L0b, with L0b confined to Ethiopia. L0a spreads widely, and even back south.

The pattern closely fits a south to north movement, but if it better fits your belief to ignore the evidence, who am I to argue?

"Well, the L3'4'6 group seems to have been specialized in coastal exploitation"

I think you're making that up. Any real evidence?

terryt said...

Sorry. I missed your earlier comments.

"I think we don't have much MSA or human presence in Southern Africa before 100,000, do we?"

But we don't know for sure that the early presence of modern mtDNA coincides with MSA culture, and vice versa. We don't know that the spread of MSA culture coincides with the spread of modern mtDNA haplogroups.

"One wonders if these last might be related to L2e".

They seem a bit too early to be realted really.

"I don't have any reason to think that slave traders massively raided the Kalahari Desert".

I doubt very much its presence there has anything to do with slavery. Perhaps very early traders moving round the Red Sea and East African coast.

Maju said...

"They're northeast of the Congo Basin?"

You did not say "northeast" but "beyond".

"Check out your own diagram. The first split is L0a'b'f'k and L0d. L0d is virtually confined to South Africa".

But not totally.

"The presence of L0d3 in Kuwait is hardly evidence for an ancient presence there".

It's an old fork: similarly old as that on L0k (Khoisan/Yemen). And there are too many L0 lineages with both regional adscriptions. And why precisely an L0d3 in Kuwait and not an L0d1'2, which is obviously much more frequent?

I don't believe in coincidences.

"L0k is also South African with a mysterious clade in Yemen"...

Like L0d3 it's half Khoisan half Arab. It's not a single mysterious lineage out of many... it's 50% of the haplogroup!

"L0f is a little further north, found in South Africa and Ethiopia (Dar-es-Salaam you say). Again with a mysterious Oman clade".

Again 50% Ethiopia/Oman (25% each) and 50% Southern Africa (in this case SE Bantu).

"I think you're making that up. Any real evidence?"

L3'4'6 centroids all fall up at Eritrea or even Yemen. What can that mean?

And we know from archaeology that some people were exploiting the coast back then at Eritrea... so why not? Makes total sense because the only "human" at Wallacea back then was H. floresiensis.

"But we don't know for sure that the early presence of modern mtDNA coincides with MSA culture"...

Well, it's a marker anyhow. But the important matter is: what's before MSA? I understand that nothing.

Correct me if I'm wrong.

"I doubt very much its presence there has anything to do with slavery. Perhaps very early traders moving round the Red Sea and East African coast".

Khoisan traders? Doesn't sound correct at all. You're forcing things again.

DocG said...

This looks really interesting Maju. I want to study it carefully first and then get back to you with my thoughts.

Maju said...

I imagined it might interest you. It's of course just my interpretation (feel free to criticize) but it's not that different from that of Behar in the reference paper. Maybe the biggest difference is that I think that there were two moments of expansion, with a period of no or limited growth between them (coalescence eras).

Another difference is my tentative proposal of L0 coalescing in East Africa (Tanganyika area maybe) instead of Southern Africa or Ethiopia. But this is admittedly tentative.

And then also, in the same line, I have "moved" the centers of dispersal of L1 and specially L2"6 slightly to the west. But that's what the composite centroid demanded. I have already mentioned elsewhere that Sudan looks like a likely center also for Y-DNA.

Finally, another difference is that I'm not considering the lineages found in West Asia or North Africa as mere product of the slave trade. Of course it may be the case with some but (with the limited evidence available) I rather suspect that it's not the case with many (some haplogroups even look North African or South Arabian in full). Further evidence might of course overrule this... or not.

Considering the number of mutations involved this represents, I suspect, about half of the history of our species (very roughly, of course).

terryt said...

"You did not say 'northeast' but 'beyond'".

I wrote, 'In fact L5 had split at 13 mut into a northeastern haplogroup (beyond the Congo Basin, L5c ...'. How can 'northeastern haplogroup (beyond the Congo Basin)' refer to any other place but northeast 'from' the Congo Basin.

"But not totally".

The only exception is Kuwait. And how come it magically appears there and nowhere in between?

"similarly old as that on L0k (Khoisan/Yemen)".

That certainly implies you're now suggesting that L0 was in Yemen and Kuwait by about 12 mutations, long before L3 had even formed, let alone left Africa. Do you really believe that's likely?

"And why precisely an L0d3 in Kuwait and not an L0d1'2, which is obviously much more frequent?"

If it was L0d1'2 you would actually have a better case for early arrival. The fact that it is little diversified from the Khoisan haplogroup is extremely indicative of post-OoA arrival. All the other haplogroups that emerged from Africa that long ago are now extremely diverse and differ greatly from their African ancestors.

"I don't believe in coincidences".

Coincidence has nothing to do with it. It's quite easy to imagine a scenario for how it happened. If the East African haplogroups in Arabia arrived later with traders thet are hardly likely to be a representative sample of East African haplogroups. Just a few would be carried out.

"Again 50% Ethiopia/Oman (25% each) and 50% Southern Africa (in this case SE Bantu)".

So? No members of either L0d3 (in Kuwait) and L0f (Oman) are found anywhere between South/East Africa and the outside-Africa regions where they are now present. Doesn't that suggest some relatively modern means of distribution?

"And we know from archaeology that some people were exploiting the coast back then at Eritrea"

But we certainly have no evidence at all that they were doing so anywhere along the South Arabian peninsular coastline. Like a fundamental Christian you may fall back on the old 'absence of evidence is not evidence of absense', but to me it severely reduces the likelihood that it was occurring.

"what's before MSA? I understand that nothing".

There must have been 'something'. Humans didn't just materialise out of thin air.

"Khoisan traders? Doesn't sound correct at all. You're forcing things again".

There you go with your brick wall mind again. We're talking female haplogroups here. Do you really believe the East African traders would be mainly women? And boy, you sure do have a short memory.
Don't you remember us discussing the origin of the Malagasay people? At that time you claimed that sea faring along the East African coast was ancient.

"I'm not considering the lineages found in West Asia or North Africa as mere product of the slave trade".

I'm sure that many North African lineages date back to the time of the OoA. L3k especially seems to be early there and other appear to get there by 26 or 27 mut. In fact I strongly suspect the OoA is very much connected to the arrival of modern haplogroups in North Africa.

Maju said...

"That certainly implies you're now suggesting that L0 was in Yemen and Kuwait by about 12 mutations, long before L3 had even formed, let alone left Africa. Do you really believe that's likely?"

I don't mean to imply that. I don't mean they were in Arabia so early. But that if they were in East Africa (between Tanzania and Ethiopia/Eritrea roughly), it's more likely that they made it to Arabia in time than if they were in South Africa, which seems too far away.

This happens with many L0 lineages but not with L0d1'2, which is the main Southern African lineage (but it does with L0d3, L0f and L0k, all shared between Southern Africa and the Red Sea area).

My reconstruction is:

1. Early L0 history in East Africa:
- t=9 - L0 at Tanganyika
- t=11 - L0a'b'f'k at Serengeti
- t=16 - L0a'b'f at Kenya
- t=19 - L0a'b at SE Ethiopia
- t=19 - L0d at Malawi
- t=20 - L0f at coastal Tanzania

2. First proto-Khoisan arrival at SA:
- t=21 - L0d1'2 at Botswana

3. Within Out of Africa
- t=24 - L0f2 at Yemen (or elsewhere at the Gulf of Aden)

4. "Young" L0 lineages coalesce at East Africa:
- t=31 - L0k at Tanganyika (one branch goes north another goes south)
- t=32 - L0d3 at Uganda

5. "Young" L0 lineages scatter to South and North:
- t=37 - L0k1 at South Africa
- t=38 - L0f1 at South Africa
- t=39 - L0k2 at Yemen

The presence of L0f and L0k lineages at Yemen and South Africa also makes me think of how Petraglia classifies "trans-Toba" Jawalpuram culture as very akin to South African MSA, what I relate in particular with L0f2.

Very tentative, I know, but that's what I get.

I can send you the centroid reconstruction map for this haplogroup: it's all very apparent in it (other haplos were more messy to map, as they have many more subclades).

Anyhow, you don't need to agree. But I can do little but explain you how I got to such conclusions.

Maju said...

"All the other haplogroups that emerged from Africa that long ago are now extremely diverse and differ greatly from their African ancestors".

Just in name. You can't find huge diversity if you have only sampled one or two or three individuals. Small haplogroups are always less diverse than large ones (by default).

"If the East African haplogroups in Arabia arrived later with traders thet are hardly likely to be a representative sample of East African haplogroups".

They should be representative of at least coastal East African haplogroups and not random minor lineages.

And until we have better data for Africa I can't say much more. I am well aware that a sample of some 650 individuals, many of them from the America, North Africa or Arabia, is not a good sample for a huge and highly diverse continent like Africa.

But it's what we have and what I'm working with.

"No members of either L0d3 (in Kuwait) and L0f (Oman) are found anywhere between South/East Africa and the outside-Africa regions where they are now present. Doesn't that suggest some relatively modern means of distribution?"

It suggests that the sample is very low for East Africa in particular and Africa in general. Please, count the individuals tagged as "Kenya" in Behar's sample, which is the only thing between Mozambique and Ethiopia: no Tanzanians, no Ugandans, no Somalians and just a handful of Kenyans.

I had to include the Hadza and Sandawe from other sources.

"But we certainly have no evidence at all that they were doing so anywhere along the South Arabian peninsular coastline".

Nor anywhere else East of Palestine and Eritrea... until Pakistan or India.

"Like a fundamental Christian you may fall back on the old 'absence of evidence is not evidence of absense'"...

Never heard before that's a "Christian" argument. It's a valid argument in any case.

Whatever the case, we have no evidence whatsoever. Paleolithic archaeology in the Muslim World is pretty bad (with some but few honorable exceptions). I can't demonstrate beyond doubt with archaeology in hand a coastal route through Southern Arabia but neither I can a continental route through Iraq and Iran. Can you?

The only suggestive evidence is the South African MSA typological identity of the toolkit of the people at Jawalpuram. But of course, they could have taken an UFO ride... can't prove they did not.

But I don't see any indication which favors a migration through Iraq-Iran either, much less for something with South African affinity (L0).

There are a few L0 terminal lineages at Egypt, Kuwait and Iran... but the only whole haplogroups that seem to coalesce over there do it at southern Arabia (Yemen in particular). And I have not disdained the Indian or Siberian lineages which might push things: there are not any, just that.

"There must have been 'something'. Humans didn't just materialise out of thin air".

Acheulean and Olduwayan. But they are considered Homo habilis/erectus/rhodesiensis/ergaster techs.

Maju said...

"There you go with your brick wall mind again. We're talking female haplogroups here. Do you really believe the East African traders would be mainly women?"

Whatever. I consider mtDNA to represent essentially whole populations. We'll deal with the Y-DNA later on, some day.

Whatever the case, I don't make sense of any traders some 100-80,000 years ago. Trade is a post-Neolithic activity, specially long distance trade.

"And boy, you sure do have a short memory. Don't you remember us discussing the origin of the Malagasay people? At that time you claimed that sea faring along the East African coast was ancient".

Sea faring and trading are not the same thing. Anyhow, we agreed that I was probably wrong about the Malagasy mtDNA issue.

"I'm sure that many North African lineages date back to the time of the OoA. L3k especially seems to be early there and other appear to get there by 26 or 27 mut."

I agree with this above. But not too much with this below:

"In fact I strongly suspect the OoA is very much connected to the arrival of modern haplogroups in North Africa".

They are connected but not necesarily and, IMO, not probably with the OoA happening through North Africa and the Fertile Crescent.

It's possible but the evidence I'm finding at the mtDNA level rather suggests South Arabia.

I happen to get way too many haplogroups coalescing (apparently) in Yemen and none in the Levant. While there are some L3 sublineages that appear quite neatly North African L3k, L3b1b, L3d1c, all are from Lybia to the West and none looks Palestine-Syria-etc. oriented. Instead we have the centroid of L3i at Yemen only 2 mutations downstream of L3 (i.e. slightly older than M). Plus the three L0 lineages also centered in Yemen or southern Saudi Arabia and L4a centered right on the Red Sea.

Would it be one lineage, would we have some other lineages centered at Jordan or whatever... then I'd be in greater doubt. But the evidence, even if not 100% conclusive, it is pretty convincing in favor of the coastal route.

terryt said...

"Anyhow, you don't need to agree. But I can do little but explain you how I got to such conclusions".

I still think you have not made a very good case for L0 being anything other than Southern African in morigin (somewhere). I mean L0d is pretty well certainly a South African lineage and the first L0a'b'f'k split leads to L0k, also basically a South African lineage (disregarding its presence in Yemen for now).

"You can't find huge diversity if you have only sampled one or two or three individuals. Small haplogroups are always less diverse than large ones (by default)".

True. But you've remarked elsewhere on the Y-hap Hs in SE Asia. Surely they've arrived in SE Asia just in the last 3-4000 years, yet already they have become distinct from their closest Indian relatives. In fact in some cases those relatives have not even been found at all. Yet here we have in Yemen haplogroups still present in Southern and Eastern Africa. Are they evidence for an early Red Sea crossing? I strongly suspect that if L4b1, for example, had arrived in Yemen at the 23 mutation level it would be much more diverse there today than it is. And perhaps it arrived in Yemen from the north via the eastern Red Sea coast.

"They should be representative of at least coastal East African haplogroups and not random minor lineages".

Not necessarily so at all. Surely we would not expect huge numbers to have moved out with any visiting sailors. Even the European haplogroups in modern America are hardly a representative cross-section of current European haplogroups, even just the coastal ones.

"but neither I can a continental route through Iraq and Iran. Can you?"

I've become very interested in mtDNA haplogroup M1. It's found in Africa but almost certainly an immigrant from SW Asia. But how long has it been present in SW Asia? We also have the interesting mtDNA N haplogroups: N1 and N2. And the Y-haps G and IJ. All could be very ancient in SW Asia. You've done a great job on the mtDNA L haplogroups. Do you have enough data, and time, to do something similar with mtDNA N? or R? M would be particularly interesting.

"much less for something with South African affinity (L0)".

I'd guess that L0 has a coastal origin in SW Asia. The only question is, how long ago did it arrive there?

"I don't make sense of any traders some 100-80,000 years ago".

You're being deliberately dense again Maju. Where have I made any such claim? And I don't make sense of L0, L4, L5 or L6 outside Africa anywhere near that long ago. Once you start claiming they were it's a short step to agreeing with German. It becomes quite easy to place L3's origin outside Africa and simply claim that most L3's re-entered Africa, and just two lineages headed east.

"we agreed that I was probably wrong about the Malagasy mtDNA issue".

It took a long time though. And you're probably wrong about the mtDNA lineages labeled L being ancient outside Africa. They probably go back no further than around the time of Austronesian arrival in Madagascar.

"all are from Lybia to the West and none looks Palestine-Syria-etc. oriented".

But we must consider the series of ancient migrations into Africa that have occurred along the North African coastline. In fact aren't those particular migrations one of your specialities? I always take any comments you make about them very seriously. So we have the possibility of many North African haplogroups being drifted out, especially when we consider the marginal conditions for existence at times over the period.

"Plus the three L0 lineages also centered in Yemen or southern Saudi Arabia and L4a centered right on the Red Sea".

I still have a huge problem placing them in the region at that time, for the reasons outlined above.

Maju said...

"I still think you have not made a very good case for L0 being anything other than Southern African in morigin (somewhere). I mean L0d is pretty well certainly a South African lineage and the first L0a'b'f'k split leads to L0k, also basically a South African lineage (disregarding its presence in Yemen for now)".

It's an alternative interpretation of the data. However you don't explain L0f, which is also split between north and south but branches out later

An "Ethiopian" origin is a third possibility. But I happen to like the Tanzanian origin the most.

You were above suggesting that the various haplogroups should have got less distant homelands, right? If so, you should favor the Ethiopian hypothesis and not the South African one.

"I strongly suspect that if L4b1, for example, had arrived in Yemen at the 23 mutation level it would be much more diverse there today than it is".

How many people do you think that could support Yemen in the Paleolithic?

I find a lot of "Yemeni" (and Omani) L haplogroups, that may be the diversity you're asking for anyhow.

"Surely we would not expect huge numbers to have moved out with any visiting sailors. Even the European haplogroups in modern America are hardly a representative cross-section of current European haplogroups, even just the coastal ones".

Are you sure? I think R1b (and to lesser extent I, R1a, J2, E1b1b1 and G2a) rule the landscape over there.

However mtDNA is different because in many regions these are essentially native (or to a much lesser extent African).

These were not traders anyhow. Traders were the ones who went to Africa, India or Indonesia. And usually what you get back in Europe is just a few erratics and not too distant of what should be expected. For example only two L lineages are in this study found in Portugal and only one in the Netherlands. Compare with the ones in Yemen please.

Can't be mere trade. Yemeni (Hadramawt) mtDNA is 35% L(xM,N) and the apportion is 9-15% in other Arab countries of West Asia (ref). Even Kurds have a 4% of such lineages!

For comparison, Ethiopians are only 55% L and Somalians 70%.

We are talking of a major component that, IMO, can't be solely attributed to "trade" (be it slave trade or whatever else).

Maju said...

"I've become very interested in mtDNA haplogroup M1".

According to your logic, it's the product of mere "trade". :P

"But how long has it been present in SW Asia?"

IMO since c. 50 Ka ago, about the same as R0.

"Do you have enough data, and time, to do something similar with mtDNA N? or R? M would be particularly interesting."

I have already dealt with these haplogroups in a most extensive manner. The mapping methods were somewhat different though, as I used diversity instead of composite centroids.

Anyhow, you can do it yourself. If you want to, you should in fact.

"I'd guess that L0 has a coastal origin in SW Asia. The only question is, how long ago did it arrive there?"

The CR mutation count clearly say that some haplogroups have been there since the OoA. My estimate and my methods anyhow.

"You're being deliberately dense again Maju. Where have I made any such claim?"

You are talking of "traders" with haplogroups that seem to be there since the OoA or just slightly after.

So who is being "dense" here?

"And I don't make sense of L0, L4, L5 or L6 outside Africa anywhere near that long ago".

I do.

"Once you start claiming they were it's a short step to agreeing with German".

You are being evil. I know where the root of the tree is.

"It becomes quite easy to place L3's origin outside Africa and simply claim that most L3's re-entered Africa".

I'm not claiming that because I have corrected my centroids according to basal diversity and common sense. But even if L3 would have originated in Yemen, from there to an American origin there's a very long distance.

There's not enough basal diversity in Arabia to make such claim: it is in Africa.

But for some lower level lineages an Arabian (mostly Yemeni) origin is very likely.

"It took a long time though".

But we agreed, so don't start throwing on me my acknowledged (apparent) errors. That's not fair play.

"They probably go back no further than around the time of Austronesian arrival in Madagascar".

That simply makes no sense whatsoever.

"But we must consider the series of ancient migrations into Africa that have occurred along the North African coastline. (...) So we have the possibility of many North African haplogroups being drifted out, especially when we consider the marginal conditions for existence at times over the period".

I'm considering sets (haplogroups) and getting centroids out of such sets. I get such centroids in Yemen and southern Arabia but I do not in Egypt or Jordan. There are diverse lineages but the centroids of their sets are elsewhere (in Lybia for instance... or in southern Arabia... or in Sudan).

So the lineages were not totally drifted out (in fact they are still some 9-15%) but no haplogroup shows signal of expansion from there. Instead they do from Yemen.

terryt said...

"However you don't explain L0f, which is also split between north and south but branches out later"

Exactly. Later, and further north. And L0a even later and even further north. And L0b later still and exclusively Ethiopian. Surely the pattern is significant.

"How many people do you think that could support Yemen in the Paleolithic?"

You seem to believe heaps of them. You have haplogroups arriving in ancient times and persisting until today. And you have Yemen providing sufficient resources for humans to survive as they moved east along the coast. So there should have been no trouble for L4b1 to have diversified considerably.

"We are talking of a major component that, IMO, can't be solely attributed to 'trade' (be it slave trade or whatever else)".

I don't think 'trade' as such. Migration surely, but the L haplogroup presence outside Africa is hardly, in itself, evidence of ancient presence.

"That simply makes no sense whatsoever".

Each individual haplogroup has limited presence, and certainly limited diversity, outside Africa. Surely that argues for relatively recent presence.

"There are diverse lineages but the centroids of their sets are elsewhere (in Lybia for instance... or in southern Arabia... or in Sudan)".

As they would be if drifted out in Northeast Africa to any extent at all.

Maju said...

"You have haplogroups arriving in ancient times and persisting until today".

I don't. They do. I just check the data and find them. I'm just the scribe, not the pharaoh.

"And you have Yemen providing sufficient resources for humans to survive as they moved east along the coast".

There are some resources: the sea and some stuff in the land. But it's not Africa nor India nor China not even Sundaland.

"So there should have been no trouble for L4b1 to have diversified considerably".

So, according to that the lineages found in Australia are, most of them, also recent because they "should have diversified considerably", and they did not. WTF!

Keep a sense of proportion.

"I don't think 'trade' as such".

That's the term you have been using so far. So don't call me dense when you can't even explain your own thoughts.

"but the L haplogroup presence outside Africa is hardly, in itself, evidence of ancient presence".

How many times I have to repeat the same: they look ancient by phylogeny and CRM count. And they are lots.

"Each individual haplogroup has limited presence, and certainly limited diversity, outside Africa. Surely that argues for relatively recent presence".

You can repeat all the time that silly "argument" but those haplogroups are at the same phylogenetic level as M or R (some even higher level). They are fucking old!

"As they would be if drifted out in Northeast Africa to any extent at all".

If we'd have such old sets in the Levant, we'd find centroids there because they would be distant from those of North Africa. We don't.

Instead even haplogroups shared between Arabia and Africa have centroids in Arabia.

But whatever. You're a dogmatic bore and that's all.

German Dziebel said...

"And I don't make sense of L0, L4, L5 or L6 outside Africa anywhere near that long ago. Once you start claiming they were it's a short step to agreeing with German. It becomes quite easy to place L3's origin outside Africa and simply claim that most L3's re-entered Africa, and just two lineages headed east."

"I'm not claiming that because I have corrected my centroids according to basal diversity and common sense. But even if L3 would have originated in Yemen, from there to an American origin there's a very long distance."

"I've become very interested in mtDNA haplogroup M1. It's found in Africa but almost certainly an immigrant from SW Asia. But how long has it been present in SW Asia? We also have the interesting mtDNA N haplogroups: N1 and N2."

It's amusing to see two firm believers in "out of Africa" to radically disagree on what constitutes the proof that African lineages ever left Africa. Luis thinks he found the "trail." Terry is afraid that by finding the trail we may find the tracks going the opposite way. So, he tries to cover it up. It's also amusing to see the same two men staring at such extra-African lineages as M1 and N1 entering Africa after a long journey from East Asia (FYI: M1 and L6 are both found in the Yemeni) and not even trying to derive L lineages from them. The fact that African L lineages aren't found anywhere past West Asia suggests that, even if they did leave Africa, they didn't go any further. Never. It's so simple. Then, M and N lineages aren't found in Africa (unless they are M1, N1, U6, which everybody agrees entered Africa), which suggests they didn't originate there. Not surprisingly they expand in East Asia, which is a far cry from East Africa. It's so simple.

So, by any rule of logic, all those L lineages found in West Asia must be remnants of the original migration INTO Africa. The fact that there are more L0, L1, L2 lineages in Africa than outside of it only means that, after a bottleneck, they re-expanded as people colonized the new huge continent. Again, it's so simple.

What kind of reasoning does either of you use to substantiate your belief in the primordiality of L lineages and their special significance to the humankind? Maybe, instead of head-butting with each other all over the web, you could try to articulate an answer to this fundamental question?

Maju said...

"What kind of reasoning does either of you use to substantiate your belief in the primordiality of L lineages and their special significance to the humankind? Maybe, instead of head-butting with each other all over the web, you could try to articulate an answer to this fundamental question?".

Sure:

All people in the world, except those within L0, belong to a set defined by mutations at these loci: 146 182 4312 10664 10915 11914 13276 16230.

That is L1''6. Only L0 is different:

In turn all L0 people have these: 263! 1048 3516A 5442 6185 9042 9347 10589 12007 12720.

There's a "Sesame Street quality" section at Wikipedia that explains that kind of stuff in terms that I'd say even little children could understand that.

Of course there may be a few cases where a back-mutation at some of those loci complicates the matter but it's mostly no big deal and easy to spot.

"The fact that African L lineages aren't found anywhere past West Asia"...

Don't LIE to my readers of I will have to censor you.

L3 clearly made it (M and N). You are very free to not believe in whatever you want but don't manipulate the data.

Meh, with the legions of multirregionalists that even today survive in the Jurasic Park areas of reality, if that would not be true they would have informed us already.

Nothing to argue, move on. You are very free to defend your honor and nonsense at the tribunals, your own blog or with your own reflection at the mirror.

Maju said...

And Terry, please, don't mention Captain America again. I don't need him messing around and claiming (with no basis whatsoever) that the phylogenies are all upside down.

I have enough with one freaky and that's you.

German Dziebel said...

"All people in the world, except those within L0, belong to a set defined by mutations at these loci: 146 182 4312 10664 10915 11914 13276 16230.

That is L1''6. Only L0 is different:

In turn all L0 people have these: 263! 1048 3516A 5442 6185 9042 9347 10589 12007 12720."

That's it?! Sure, the majority of those L0 people speak click languages. All languages can be divided into those with clicks and those without clicks. Purely abstractly. But no serious linguist would claim that the original human language had clicks and that Bushmen are the only one who preserved them. There were mavericks such as Stopa who thought that clicks are halfway between ape and human language but it's a long-gone fallacy. It's pretty clear that clicks is a local Bushmen-specific evolution (or Sandawe-specific evolution). Same logic should work for the genes.

"Of course there may be a few cases where a back-mutation at some of those loci complicates the matter but it's mostly no big deal and easy to spot."

Easy for me. Apparently, impossible for you.

""The fact that African L lineages aren't found anywhere past West Asia"...

Don't LIE to my readers of I will have to censor you.

L3 clearly made it (M and N). You are very free to not believe in whatever you want but don't manipulate the data."

I never lie. I have two doctorates, the fact that pretty much safeguards your readers against that. M and N aren't found in Africa, unless in derived states. It's a fact. This makes them essentially non-African haplogroups. African L lineages are barely found outside of Africa. That's another fact. The "out of Africa" theory can't even live up to its own name. At least we should call it an "always in Africa" or "Africans R Us" theory. M, N and L3 are separated by a long string of unattested mutations and thousands of geographic miles. A good phylogeny should mirror phylogeography. That's a third fact for you. Where are the lies?

You can worship L0 as much as you want and you may even begin to talk in clicks for extra authenticity, but science is about proof, not belief. You can at best stitch these objective gaps with your imagination.

"Meh, with the legions of multirregionalists that even today survive in the Jurasic Park areas of reality, if that would not be true they would have informed us already."

Multiregionalists also believe in continuity in Africa. So they are in cahoots. Out of America is the only theory that questions it.

Maju said...

There are L0 people who speak click and there are L1"6 people who do. There are L0 people who speak without (mostly) any clicks and the same for L1"6.

"But no serious linguist would claim that the original human language had clicks"...

I would not be too surprised: clicks are found in most languages though seldom as consonants (but they do exist as distinct sound-signals). But it doesn't really matter because L0 is just one of two branches and languages don't matter at all.

I'm just sooo bored of people who claim absurd things based on absurd speculations on languages that I don't care the least about languages.

"I never lie".

You do. You know things that you silence and you ask questions for which you know the answer. You are just a manipulator: a pathetic manipulator without anything better to do than lurk in other peoples' blogs in the hope of getting someone who listens to your idiotic out of America nonsense, in which not even you believes.

"I have two doctorates"...

I don't care: I know just too many career people who know nothing and too many high school dropouts who know much more, have more interest in truth and are more intelligent and, specially, sincere with themselves and the rest.

"M and N ... African L".

From the viewpoint of phylogenetics M and N are L-something. There's no such thing as L anymore: L was described when phylogenetics was in its infancy. Now we know it's two basal lineages (L0 and L1"6) and that M and N are nothing but derivatives of a derivative of a derivative (L3) of a derivative (L3'4) of a derivative (L3'4'6) of a derivative (L2'3'4'6) of a derivative (L2"6) of L1"6. Phylogenetically M and N are at the level of L0d1c1 for example.

You just want to dance with obsolete nomenclature because that way you justify your freaky obsession. I don't care: I just don't want to hear of it.

"M, N and L3 are separated by a long string of unattested mutations"....

That's a LIE. M is just 3 or 4 mutations (depending on whether you count HVR or not) downstream of L3. N is five mutations. You know that perfectly but you just want to fuck everybody around with idiocies and lies like that one.

I don't need that shit. Get lost.

Maju said...

Ah, and I really hate people who hide behind titles and other medals. If you know you will show it in your discourse, if you're an ignorant you can cover your walls with titles that they won't fix it.

But if you are an ignorant and you try to cover that by claiming doctorates, PhDs or whatever... then you karmically deserve the worst of destinies: to be mocked by for arrogant and stupid, to see your titles spitted upon and you and your very soul as well.

So step down and get some "degrees" in hard work, show me the roughness of your hands and that your words have any meaning at all. Your doctorates surely have good use at the toilet.

terryt said...

"So, according to that the lineages found in Australia are, most of them, also recent because they 'should have diversified considerably', and they did not".

That statement would be far more convincing if it were true. But it's not.

Not only has Melanesia/New Guinea/Australia mtDNA haplogroup diversity been recognised and labelled, it is almost completely distinct from that outside the region and is even separated between the two major regions (apart from one or two New Guinea haplogroups found in Northern Australia, almost certainly relatively recent arrivals). This is completely unlike the mtDNA L haplogroups you claim as evidence for a remnant of the original OoA.

Within New Guinea/Melanesia mtDNA haplogroup Q has been classified as Q1, Q2 and Q3; haplogroup P as P1, P2 and P4a. Within Australia mtDNA haplogroup P has been classified as P4b, P5 and P9; S as S1, S2 and S5. In fact P4 is really the only haplogroup common to both Australia and New Guinea, but New Guinea has P4a and Australia P4b. And that's ignoring the M haplogroups distinct in each region.

Distinct. Unlike the majority of your favoured non-African mtDNA L haplogroups. And of the non-African haplogroups that are distinct most are no older than the 30 mutation level. So, think again.

Maju said...

I mean the lineages, each of them:

N13 and N14 have no known sublineages, O (N12) has only one proposed sublineage. Even S (which is the most important AA single lineage) and M42 are not really that diverse.

Together they are "diverse" but just like the various Arabian L lineages are, mutatis mutandi: a diverse bunch but not a bunch of diversified lineages.

terryt said...

"Even S (which is the most important AA single lineage) and M42 are not really that diverse".

But they're more diverse than any single non-African L lineage. And they can have emerged from Africa no earlier than your proposed ex-African L lineages.

"a diverse bunch but not a bunch of diversified lineages".

That describes far better your non-African L lineages. The Arabian L lineages are diverse because they were arrived from diverse locations, relatively recently.

Maju said...

"But they're more diverse than any single non-African L lineage".

Not sure but the difference is tiny (and Australia many times the size of Yemen or even Peninsular Arabia - and not more arid).

Of all the Australo-Melanesian mtDNA lineages the only one clearly diverse is P.

"That describes far better your non-African L lineages".

It describes well both: a diverse bunch of undiversified lineages, in Arabia as in Australia.

"The Arabian L lineages are diverse because they were arrived from diverse locations, relatively recently".

The same that all Australian lineages seem arrived from Wallacea/New Guinea (or SE Asia in general), all Yemeni lineages appear to sprout from East Africa.

The cases are fully comparable: both seem to imply not a single founder effect, product maybe of a rare single crossing, but the arrival, maybe along some lengthy period, of a relatively large number of initial settlers, carrying some diversity from their ancestral homeland beyond the sea. Diversity that was preserved to some extent till now.

Maju said...

"And they can have emerged from Africa no earlier than your proposed ex-African L lineages".

Let's see:

L4b, L3i and L0f2 are older than M and found at both sides of the sea (just like M or N but at smaller scale).

L0a1b2, at 27 mutations (older than N), is only found in Saudi Arabia.

There may be others... I'm too lazy to bother looking for all.

You just don't want to believe in that possibility but there it is.

terryt said...

"at 27 mutations (older than N)"

That's mutilating the evidence. N forms at 23 mutations. It's just that it has a long period before it expands, but there's no evidence it hadn't left Africa long before it expanded further. Interesting that you're arguing N's long tail is evidence it moved out of Africa relatively late, yet claiming L4b1's long tail is evidence for its early exit.

"L4b, L3i and L0f2 are older than M and found at both sides of the sea"

Both sides, note. So you're arguing that this provides convincing evidence for their ancient presence outside Africa. Surely it's just as possible, in fact more probable, that their presence outside Africa is post OoA.

"You just don't want to believe in that possibility but there it is".

A 'possibility'? Yes. But hardly 'definite evidence', in fact hardly 'probability'. I promise I'll let you have the last word here.

Maju said...

I hope that you are intelligent and self-critical enough to realize that you are cheating shamelessly by using blatant double standards: on one side you demand me to treat pre-N (L3* leading to N) as "N" and then you also demand that I don't do the same with the Arabian L lineages.

If you'd apply your standard #1 (the one you ask me to use with N and only with N) to L6 or most other Arabian L, you'd corner yourself.

And that's what you should do if you had the slightest remnant of self-criticism and honesty.

So... are you serious or are you a damn preacher with a bubble-over-the-head syndrome? Are you scientific or fanatic?

Maju said...

Erratum "light-bulb", not "bubble".