And your little dog, too!

As I have mentioned before,  the mtDNA of European hunter-gathers seems to be very different from that of modern Europeans. The ancient European mtDNA pool was about 80% U5b – today that lineage is typically found at 10% frequency or lower, except in northern Scandinavia. Haplogroup H, currently the most common in Europe, has never been found in early Neolithic or  pre-Neolithic Europeans.   Less has been done with ancient Y-chromosomal DNA, but the data we have suggests that the lineages that were common in the early Neolithic and pre-Neolithic are rare today.  R1b is the most common y-chromosomal haplotype today in Western Europe:  it has never been found in Neolithic or  pre-Neolithic Europeans.  We find G2 in the early Neolithic: today it is rare, except in Sardinia and some mountainous regions of Italy and Sicily. It sure looks as if we’re talking near-complete replacement – which means that the historical process involved does not look much like a peaceful, diffusion-style range expansion.  Perhaps more like the Death Song of Ragnar Lodbrok, which abounds in phrases like this: “Where the swords were whining while they sundered helmets”

Interestingly, there is a very similar  pattern in canine mtDNA.  Today Europeans dogs fall into four haplotypes: A (70%), B(16%), C (6%), and D(8%).  But back in the day, it seems that the overwhelming majority of dogs (88%)  were type C,  12% were in group A, while B and D have not been detected at all.

The ancestors of today’s Europeans didn’t fool around.

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39 Responses to And your little dog, too!

  1. dearieme says:

    Dog-eating bastards!

  2. dearieme says:

    I withdrawing that rather vigorous noun: the thought nonetheless occurs that maybe the incomers hailed from Korea?

  3. Peter Frost says:

    Almost complete population replacement? Unlikely. Among other things, it would mean that Middle Easterners came to look like northern Europeans within a very tight timeframe. Two or three thousand years, perhaps less. That’s possible, if you postulate very strong selection pressures. But I don’t see that in your model.

    Natural selection is likelier. We see high incidences of the U5 haplogroup both before and after the transition to farming from hunting/gathering/fishing. This was the conclusion of a study of 92 Danish human remains that ranged in time from the Mesolithic to the Middle Ages. There was genetic continuity from late hunter/gatherer/fishers to early farmers:

    “The extent to which early European farmers were immigrants or descendents of resident hunter-gatherers (replacement vs. cultural diffusion) has been widely debated, and new genetic elements have recently been added. A high frequency of Hg U lineages , especially U5, has been inferred for pre-Neolithic Europeans based on modern mtDNA data, with Hg U5 being fairly specific to Europe. […] Our study therefore would point to the Early Iron Age and not the Neolithic Funnel Beaker Culture as suggested by Malmstrom et al. (2009), as the time period when the mtDNA haplogroup frequency pattern, which is characteristic to the presently living population of Southern Scandinavia, emerged and remained by and large unaltered by the subsequent effects of genetic drift” (Melchior et al., 2010)

    So the sharp genetic divide was not between late hunter/fisher/gatherers and early farmers. It was between the earliest farmers and groups that had been farming for at least a millennium. The evidence points to natural selection and not to population replacement.

    Yes Virginia, mtDNA responds to natural selection. Some haplogroups seem to reflect a tradeoff between thermogenesis and ATP synthesis. The decline of U-type haplogroups may reflect differences in physical activity between them and hunter/fisher/gatherers.

    Reference

    Melchior, L., N. Lynnerup, H.R. Siegismund, T. Kivisild, J. Dissing. (2010). Genetic diversity among ancient Nordic populations, PLoS ONE, 5(7): e11898

      • Peter Frost says:

        I’d rather not. The purpose of betting is to humiliate one’s opponent. That isn’t my goal.

        But if you insist, a bet would have to meet the following conditions:

        1. Amount: $10,000, to be paid by the loser to Steve Sailer
        2. For the purposes of the bet, “near-complete replacement” means that less than 10% of the current European gene pool comes from Mesolithic Europeans.
        3. In the event of disagreement, the bet would be settled by a panel of three anthropologists: Henry Harpending, an anthropologist chosen by me, and and anthropologist chosen by you.

      • dave chamberlin says:

        It would be nice to hear this debated between the two of you. Forget about money or who wins, that will just turn the two of you into self seving salesman rather than the curious scientists that you both are. Just my two cents but science blogs have grown to such a degree that debates between scientists of opposing view points could really take off if done right. Kind of a blogging heads but purposely made confrontational by setting known spokesmen of opposing viewpoints against one another. The only real winner would be the audience.

    • gcochran9 says:

      Who said that North Europeans came from the Middle East? I sure didn’t. Second question: I did mention Y-chromosomes also turn over at about the same time. Why?

      • Peter Frost says:

        Your argument is that Europe’s transition from hunting/gathering/fishing to farming occurred through population replacement. If these incoming farmers didn’t come from the Middle East (the usual assumption), where did they come from? Africa? East Asia?

        The Y chromosome change occurred after the above transition (like the mtDNA change). It too was probably driven by natural selection.

    • jb says:

      I really would like a nice clear explanation for the difference in appearance between northern and southern Europeans. A naive explanation, one that I sometimes give my friends, is that when agriculturalists from the Middle East spread into Europe, they colonized southern Europe quickly, because it was easy to reach by sea, and diffused into northern Europe more slowly, and as a result the northern Europeans look more like the original pale skinned Paleolithic Europeans, while southern Europeans look more like the darker Middle Eastern immigrants. Is this account completely bogus in light of recent discoveries, or might there still be something to it (with various population replacements layered on top maybe)? It really is a nice story, and provides a simple and satisfying intuitive explanation for a population difference that even non-scientists are very aware of and interested in.

    • whatever says:

      They did not come from the middle east. Ancient y-dna says plains of what is now Russia, south Siberia, or even as far as Baikal lake and central Asia. Plenty of time and climate conditions to become what they became, before they arrive in Europe and eat all dogs. As for the mt dna in Europe, it is from everywhere; some of it came from paleolithic hunter gatherers, some of it is a left over from the early neolithic farmers and some followed the y chromosomes of the pastoral nomads from the south Siberian plains (i.e. the people, who gave the rise to the most of contemporary European population) – there are matches b/n both contemporary mt dna and contemporary y-chromosomal dna and ancient dna, extracted from the Siberian kurgans. See for example Keyser et al. 2009, but many others too – it aligns with the finds of D. Anthony.

      • uh says:

        Don’t pay attention to that huge R1a swathe cutting through Eastern Europe and Asia. Has nothing to do with the Aryans; they were brown folk from Anatolia.

        lolz

    • It was between the earliest farmers and groups that had been farming for at least a millennium. The evidence points to natural selection and not to population replacement.

      In recorded history, there have been numerous incidents of near total replacement of farmers by farmers – the classic Greeks were totally replaced by today’s “Greeks”, the Britons were totally replaced by the Anglo Saxons. We are a genocidal species. Humans are chimps that have specialized for war.

    • harpend says:

      Good choice, Peter. I stand ready to provide the fairest judgment that money can buy.

    • gcochran says:

      I was thinking more of a dollar, rather than ten grand. I would only make a bet like that If I was royally pissed at you: I’m not. But I won’t even bet a a dollar unless I’m damn sure.
      Everyone working on ancient DNA (for example the Geographic Project) is talking about finding indications of _multiple_ layers of replacement in Europe. David Reich and Patterson also: we should see a paper soon. Everything published agrees with this. Moreover, although I would be the last to deny the possibility and importance of selection, most loci are _not_ under strong selection. Selection making these big changes in mtDNA frequencies – big enough that the Fst in mtDNA between old European hunters and modern Europeans is greater than that between Australian Aborigines and modern Europeans – I could at least imagine believing. With difficulty. Suggesting that it happened in Y-chromosomes too – that’s a stretch. I don’t believe it. Moreover, I look at ancestry estimates (Admixture, Frappe, etc), and it’s clear that there was a southern European replacement out of the Mideast (with Sardinians as as a relatively pure example), and, at the same time, that this was followed by at least one _other_ wave of replacement, primarily in northern Europe. The logical bet is Indo-Europeans, out of somewhere in Russia, likely the Pontic-Caspian steppe. If you are worried that there wasn’t time for northern Europeans to develop all the odd hair and eye colors, stop worrying: the color variants were already there in the Indo-European urheimat. Certainly they were there in Andronovo culture, circa the Bronze Age: they’ve been found, in ancient DNA. But if you look at Uighurs or the Kalash, you already knew that those color variants extended deep into Asia.

      • Sean says:

        Maybe there was virtually total replacement in the male line, and the male hunter gatherers’ Y-chromosomes lost out big time. (though more than in any similar conquest I’m aware of). “Selection making these big changes in mtDNA frequencies …I could at least imagine believing. With difficulty.” It seems to me selection on mtDNA is certain, else what happened to the European HG women. Indo-Europeans’ expansion would be spearheaded by bands of young men. Unless the title of this post is hinting at a mass “A Boy And His Dog ” type scenario, there would be offspring, and their mtDNA would be from the female line; it would not change. How then could it disappear?

        There are political implications to Europeans’ being the product of genetic replacement and to become known as a scientist who thinks Europeans HGs were not replaced may have personal costs attached. (ie career implications) I dare say some scientists may be keeping quiet. RE. “odd hair and eye colors”. Women color their hair. They know what ‘odd’ hair color is for.

  4. dave chamberlin says:

    Another parallel between pooches and humans is the ever changing argument of where they come from. One theory proposes all domestic dogs came from China, another one that they all came from the middle east, and now it appears Fido’s ancestors jumped the fence and got frisky with the locals, at least enough to create the phenomenal diversity in dogs we see today. The totally out of China theory for dogs is as wrong as the totally out of Africa theory is for humans.

  5. Sean says:

    They killed the men, What did they do to the women ? As I understand it, we inherit our mtDNA from our mother only .

    Peter said:” Some haplogroups seem to reflect a tradeoff between thermogenesis and ATP…”.
    Northern Scandinavia is cold.

  6. Sean says:

    Mother’s curse: the effect of mtDNA on individual fitness and population viability..”Ironically, deleterious mtDNA mutations that affect only males, such as those that impair sperm function, will not be subject to natural selection because mitochondria are generally maternally inherited. Thus, such mutations may reach high frequencies affecting the viability of natural populations.”

    When European hunter gatherers adapted agriculture the MEN with the crap , ie thermogenisis adapted mtDNA (Read Nick Lane)- were suddenly at a disadvantage and got out-bred in the resultant population explosion. I dare say a lot of their Y-chromosomes disappeared too for similar reasons ( ie non competitive in the new fitness environment). The winners were the few aboriginal European hunter gatherer men with mtDNA and or Y chromosomes that made them able to father many children. Foreign interlopers’ mtDNA and especially Y-chromosomes DNAalso did well of course.

    The issue of different mtDNA being selected for with the coming of agriculture Women with mitochondrial haplogroup N9a are protected against metabolic syndrome

    • rob says:

      To look for selection on mitochondrial genes, wouldn’t it be easier to look at nuclear genes that code mitochondrial proteins? Then we can use linkage disequilibrium etc, and also ignore selection pressures that are peculiar to mtDNA? If the nuclear genes are rarely under positive selection, then it’s not very likely that selective pressure on mtDNA could cause a variant to sweep the population.

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  8. Matt says:

    http://dienekes.blogspot.com/2011/02/human-migration-and-cultural-change-in.html

    “Another point of interest is that the Dnieper-Donetz samples from Dereivka and Aleksandrija have close parallels to the Portuguese Mesolithic! This tends to reinforce the view that has emerged from the study of mtDNA of a fairly homogeneous pre-farming substratum that stretched from the Atlantic well to the east.

    That does seem to push out the limit quite far to how far East and Central a non-Middle Eastern European replacement population would have to be from, to avoid being part of the same group (and thus have different mtDNA and y-Dna). And push it further east (and south, further north seeming improbable?) than the hypothesised Kurgan homeland.

    And if they were from an area with this position, how relatively shifted towards South and East Asians would we expect them to be, compared to Middle Eastern populations that (presumably?) have not seen such replacement?

  9. Peter Frost says:

    “Everything published agrees with this.”

    Then you and I live in parallel universes.

    I recognize that there is much debate on this subject. At one end, some people argue that Mesolithic Europeans are ancestral to over 80% of the current European gene pool. At the other end, some people say the figure is only 25 to 30%. But nobody, except you, is arguing for the hypothesis of near-complete population replacement.

    At present, the majority opinion tends toward the first viewpoint, i.e, Middle Easterners initially established pioneer farming settlements in central Europe but were over time largely replaced by native farmers. Rowley-Conwy (2011, p. S434) describes this new model:

    “Our explanations must now rest on two major foundations: most Neolithic genes were native, but the major domesticates were exotic. Small-scale rather than continent-wide migrations are the best way to integrate these into one model. Agriculture in a region may have been introduced by immigrants, but that does not mean that the immigrants carried mainly Near Eastern genes (Richards 2003; Rowley-Conwy 2004b; Zvelebil 2005). The LBK, for example, originated in the Carpathian Basin; the population that moved westward emerged there carrying a complex mix of European and Near Eastern mtDNA and no doubt picking up more as it moved.”

    You express skepticism that natural selection could have produced so much change in mtDNA frequencies “between old European hunters and modern Europeans.” Actually, as I pointed out, the change was between the earliest farmers and somewhat later farmers who apparently shared the same cultural tradition. So, yes, natural selection can produce such changes over that kind of time frame. It’s not clear to me why this strikes you as impossible, but in any case the question is moot. It happened – quickly and among people of the same culture.

    I could go on. The spread of Indo-European languages was not a matter of population replacement, at least not primarily. Germanic languages show strong evidence of a Finnic substrate. Teach a Finn to speak Indo-European and you get a German. There is much evidence of conquest in European prehistory, and of the conquered emulating the conquerors, but we see little evidence of ethnic cleansing.

    Reference

    Rowley-Conwy, P. (2011). Westward Ho! The Spread of Agriculturalism from Central Europe to the Atlantic, Current Anthropology, 52 (S4), S431-S451

    • gcochran says:

      He’s leaning on Soares et al. Their argument is that if an mtDNA or Y-chromosome haplogroup’s estimated time of origin was, say, 15,000 years ago, it must have originated in Europe rather than being a Neolithic or post-Neolithic import.

      That’s obviously wrong. If a fair number of people migrate in, rather than a single individual, lots of variation is preserved, and the TMRCA in the immigrant group is essentially the same as in a population that stayed in place. Look, if you look at R1b in Ohio, it looks just as old as it does in western Europe: does that mean that whites have been living in Ohio for 15000 years or whatever?

      Basically, Soares and company _assume_ that there were never any large-scale migrations into Europe,and then use that to prove that there were never any large-scale migrations into Europe.

      I, on the other hand, am relying on the observed data: most of the currently common mtDNA haplotypes in Europeans have never been seen at all in ancient DNA from European hunter-gatherers. Same of Y-haplotypes. And assuming massive selective changes in both mtDNA and Y-chromosomes – which you are doing – is way more unlikely than assuming such change in mtDNA – itself already unlikely.

  10. gcochran says:

    Rowley-Conwy is relying on outdated and obviously wrong genetic analyses. And through him, you are as well.

  11. Peter Frost says:

    “Outdated and obviously wrong” … Uh, the Rowley-Conwy article was published last year in Current Anthropology, which is the top journal in the field.

    You started off with a falsifiable hypothesis, i.e., the transition to farming in Europe was a demographic change — one population was replaced with another . I falsified your hypothesis by pointing out that the genetic change happened after the transition. In other words, the genetic divide was not between late hunters/gatherers/fishers and early farmers. It was between early farmers and not-so-early farmers.

    You’ve now shifted the population replacement to an unspecified point later in time and to an unspecified cause. I have no way of proving you wrong. Your hypothesis is unfalsifiable.

    Your comparison to White Americans in Ohio is thus beside the point. We have historical records linking those people to ancestors in the British Isles and elsewhere in Europe. If need be, we could ask those Ohio residents where their ancestors came from.

    This doesn’t hold true with your hypothesis. You’re arguing that present-day Europeans are descended from a population (or populations) that entered Europe at an unspecified time and from an unspecified place. Finally, this postulated non-European population no longer exists, and there are no records of its existence.

    I can’t argue with that kind of argument. It’s unfalsifiable.

  12. gcochran says:

    Peter, Ronwy-Conwy’s paper relies on Soares paper, “The Archaeogenetics of Europe”, which came out on February 22 2010. This misses a lot of recent work on ancient DNA: it’s a fast-moving field. The Soares paper dismissed the recently developed ancient mtDNA evidence, which shows almost complete disjunction between hunter-gatherer mtDNA in Europe and modern Europeans. it also shows near-complete disjunction between those hunter-gathers and early LBK farmers, and a not quite as large, but still very large, difference between those LBK farmers and current Europeans. I would guess that Soares et al dismiss it because the ancient DNA data shows that all the results of their research program – which involves trying to estimate past locations and origins of different mtDNA haplogroups – is completely wrong.

  13. Halvorson says:

    I can’t figure out if I’m much dumber than Peter Frost or much smarter: isn’t Greg insinuating that that Mediterranean farmers replaced the native HGs and then were themselves largely replaced by Indo-European horselords from the steppe?

    • Sean says:

      So all the aboriginal European hunter gatherers, including the women, were completely killed off by ME agriculturists. And then the ME agriculturists, women included, were in turn killed off by Indo-Europeans. Those fellows had a lot to learn about conquest I think

      • Halvorson says:

        Well, mtDNA U still exists at a frequency of around 10 percent in modern Europe (max is around 20 in Finland I think), so even in a scenario with no negative selection the women weren’t completely replaced. And they wouldn’t have to have been exterminated, just massively outnumbered by incoming farmers who could produce 10-100 times as much food using the same amount of land. But like Peter mentioned, there’s a still a discontinuity between early and late Neolithic mtDNA. I still don’t quite buy the idea that Indo-European pastoralists would be able to so completely overwhelm long established farmers even on the maternal line, but I think that’s what C+H are getting at.

      • Sean says:

        As i read it what C&H were getting at in the 10,000 Y E was not replacement, but that the Indo-Europeans, the Kurgans, practiced elite dominance rather than replacement. Now, it seems to me, we are being told that the Indo-Europeans brought blonde hair and blue eyes (and maybe white skin ) to Europe from the Pontic-Caspian steppe. Despite the fact that the Kurgans were the ones with least need for antirachitic alleles (which white skin is not) as the Kurgans had had milk.

  14. Ortu Kan says:

    And here’s the picture from New World dog mtDNA:

    “The few sequences from breeds of New World origin (the Eskimo dog, Mexican hairless, Alaskan husky, Newfoundland, and Chesapeake Bay retriever) and from Oceania (the Australian dingo and the New Guinea singing dog) were indistinguishable from those of Eurasian dogs (5).”

    “Six of 12 ancient Latin American haplotypes are grouped in clade a and include sequences found in dog remains from Bolivia, Peru, and Mexico (Fig. 1). No sequences from clade a have been found in samples from over 350 modern dogs (17). The upper bound of a 95% confidence limit for the frequency that sequences from clade a could have in this modern sample and be missed (an observed frequency of zero) is 1.0% (24). Consequently, the absence of clade a sequences from modern dogs suggests an extensive replacement of native American dogs by those introduced by Europeans. These lineages could be surviving in some unsurveyed modern Native American breeds or local dog populations (14,25). However, genetic analysis of a diverse sample of 19 Mexican hairless dogs (xoloitzcunitle), a distinct ancient breed that has been present in Mexico for over 2000 years (25), only revealed mtDNA sequences previously observed in dogs of Eurasian origin (26). The absence of ancient North and South American dog haplotypes from a large diversity of modern breeds, including the Mexican hairless, illustrates the considerable impact that invading Europeans had on native cultures.”

    • Ortu Kan says:

      Their a is a subclade of the A mentioned earlier, outside of which (in B) only a single ancient American haplotype fell — one of three from Tula (Hildago), Mexico, 1400 yr B.P.

  15. Joe Walker says:

    Quote: “So all the aboriginal European hunter gatherers, including the women, were completely killed off by ME agriculturists. And then the ME agriculturists, women included, were in turn killed off by Indo-Europeans.”

    Isn’t it possible that the newcomers brought diseases with them that helped to kill off the pre-existing European populations?

  16. MAR says:

    Speaking of dogs, does Bryan Sykes still have a dog in this fight? I know that at one time he opposed the idea of population replacement of European hunter-gatherers. Is he still pursuing this line? Will he publish more on it?

    Given the Ub5 divide between Southern and Northern Europe, isn’t is possible that ME farmers settled in Southern Europe but that agriculture spread culturally to Northern Europe? Might there have been more replacement in the South but little in the North?

  17. MAR says:

    “There are political implications to Europeans’ being the product of genetic replacement and to become known as a scientist who thinks Europeans HGs were not replaced may have personal costs attached.”

    In some respects, a larger picture teased out from GC’s thesis might be messier (if I’m reading him correctly) by filling in some blanks: HGs wiped out Neanderthals (albeit with some admixture), ME farmers wiped out HGs, IEs from Russia wiped out MEs. That’s bloody.

    • whatever says:

      No surprise here; for very long time, Europe has been – and still is – the grave yard of nations; like no other place on earth; numbers have left their bones there; few have survived.
      The reasons are diverse; many of them – environmental.

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