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14 – And Moses was wroth with the officers of the host, with the captains over thousands, and captains over hundreds, which came from the battle.

15 – And Moses said unto them, Have ye saved all the women alive?

16 – Behold, these caused the children of Israel, through the counsel of Balaam, to commit trespass against the Lord in the matter of Peor, and there was a plague among the congregation of the Lord.

17 – Now therefore kill every male among the little ones, and kill every woman that hath known man by lying with him.

18 – But all the women children, that have not known a man by lying with him, keep alive for yourselves.

- King James Bible, Numbers 31

In the 20th century the Lithuanian archaeologist Marija Gimbutas posited that the emergence of pre-Christian European culture went through two phases after the Mesolithic. First, there were the Neolithic Old Europeans who brought agriculture. Then there were the Kurgan people from the steppe, who brought Indo-European languages and warlike patriarchal values to the continent.By the 1990s many archaeologists had turned against the Kurgan model of Indo-Europeanization, leaning rather toward the proposition that the Old Europeans themselves were Indo-Europeans. I believe that the latest work in genetics, utilizing powerful statistical inference techniques leveraging genomics and computational biology, and ancient DNA, suggest that Gimbutas was right in terms of the role of the Kurgan people as promoters of Indo-European culture in Northern Europe. Even those who supported the Kurgan hypothesis, such as David Anthony, were apparently shocked at the magnitude of the genetic turnover.

warbefore But Gimbutas probably went very wrong is the idea that Old Europeans were a peaceful and matriarchal society. First, though there are matrilineal societies, and matrifocal societies, to my knowledge there are no matriachal societies which are analogs to the patriarchies you might find in the modern Arab world or ancient Athens (and frankly, most agricultural and post-agricultural societies). Certainly there were societies where powerful women were shaping the course of events. This influence may even be institutionalized (I’m thinking of the Iroquois as an instance of a case). But there were no societies where rulers were exclusively women and men were forced into roles of total passivity in matters of war and politics, and property as a class.

That’s the truism as informed by what we know from surveying cultures in the historical record and extant today. But there is a spectrum of empirical phenomena in terms of magnitude. During the Roman Empire the women of the Latin West continued to have liberties and freedoms that were customary for them during antiquity (the power of the Julio-Claudian women and Theodora seem less shocking when considering the public prominence of elite women during the Republican period, which some ascribe to the role of Etruscan women in their society). When the focus of Roman power shifted toward Constantinople in the 4th century, one visible marker distinguishing elite women of western cultural affiliation, as opposed to those who were of the Greek nobility, is that the latter were often veiled, perhaps echoing the seclusion of ancient Athenian women of good family.

Similarly, though Japanese civilization is influenced, perhaps even derived, in large part from Chinese civilization, one major distinction between the two is that the in the ideal and often in practice the Chinese have subordinated military values to civilian ones to an exceptional extent for a pre-modern society. In contrast, the Japanese developed a military aristocracy which eventually superseded the civilian nobility. This results in the anachronistic romanticization of a martial ethos such as in bushido, which has no clear analogy in the Chinese world view. Obviously here I am not saying that the Chinese were a purely pacific people. And there were ages when martial values were ascendant, for example the early Tang. But the fact that the founder of the Song dynasty, a general, encouraged a demilitarization of his ruling class makes much more sense in light of the ethos of Chinese elite culture going back to the end of the Warring States period. In contrast, the Western aristocracy, often directly descended from Germanic warlords, have retained an ethos where physical violence and competition is more meritorious. The emergence of firearms necessitated a shift away from direct front-line combat to minimize casualties, and a channeling of energies into patronage of high culture and foppish self-cultivation. But even today the princes of the House of Windsor continue to serve in military professions, putting the role of the soldier in Western society in stark relief as one of esteem.

51PS1EGohbL._SX309_BO1,204,203,200_ I bring this up to reiterate that though we see the past through a dark mirror, we must filter its probabilities through what we know of societies today, and those that are historically attested. Human phenomena is not infinitely flexible, but exhibits modal peaks across the distribution of possibilities. Our expectations should not be uniform and agnostic. The Old Europeans may have been gynocentric pacifists, but if they were then they were sui generis among human societies. As time machines are not feasible we will never truly know in a direct sense what they were like. Rather, we must look to aligning material remains with theoretical expectations given what we know about the nature of human societies. Interpretation will always occur. The key is to obtain the proper framework to generate true inferences. In Lawrence Keeley’s War Before Civilization the author observes how the objects which might be useful as weapons in graves have often been interpreted as “ritual” markers of status, as if conspicuous consumption was always the primary form of status competition. Written in the 1990 War Before Civilization was a seminal work taking on the neo-Rousseauan model head-on, that war was somehow a contingent invention of civilization. A terrible mistake.

A recent paper in PNAS puts the final nail in the coffin of this strong form of the neo-Roussseauan paradigm, which now has little support even from scholars such as Brian Ferguson. The paper is The massacre mass grave of Schöneck-Kilianstädten reveals new insights into collective violence in Early Neolithic Central Europe:

The Early Neolithic massacre-related mass grave of Schöneck-Kilianstädten presented here provides new data and insights for the ongoing discussions of prehistoric warfare in Central Europe. Although several characteristics gleaned from the analysis of the human skeletal remains support and strengthen previous hypotheses based on the few known massacre sites of this time, a pattern of intentional mutilation of violence victims identified here is of special significance. Adding another key site to the evidence for Early Neolithic warfare generally allows more robust and reliable reconstructions of the possible reasons for the extent and frequency of outbreaks of lethal mass violence and the general impact these events had on shaping the further development of the Central European Neolithic.

The body of of the text engages in a deep osteological analysis, but in the language of the street, “they fucked these people up.” In particular, the victims seem to have had their lower extremities maimed or crushed. If they were still alive when this occurred then it was clearly a form of torture. If they were dead, then it was clearly a spiteful mutilation of the dead, and the valence has to be symbolic rather than utilitarian. The victims in the assemblage exhibited a curious demographic pattern. There were infants below one year of age, as well as young children, but no older children or adolescents. The only two adult women were over the age of forty. The rest of the adults killed were men.

We can’t know what happened with certainty. These were preliterate people. But with what we know about the nature of human culture it seems that an obvious narrative presents itself. As noted in the paper this was an LBK site. But, it seems that the community was on the border of two LBK trade networks (as inferred from the distribution and character of material remains). On the frontier of agricultural production, when land is in surplus, one can imagine that there was little inter-group conflict between LBK coalitions. What we would probably term “tribes.” Additionally, there was almost certainly a “meta-ethnic frontier” which Mesolithic hunter-gatherers, who we now know were genetically and physically very distinct from the LBK people (naively projecting genetic variance statistics, their difference was in the ballpark as that between modern Chinese and Northern Europeans, Fst ~ 0.10).

But what happens when Malthusian constraints begin to close in? In the Moral Consequences of Economic Growth Benjamin Friedman suggests that in American history economic stagnation and stress lead to greater xenophobia, and reduced openness. And one doesn’t need a deep history lesson to observe what occurred in Europe during the 1930s. Retrenchment invariably leads to turning back to collective units of organization and protection. Once the LBK reached a stationary state, which reduced marginal returns to labor input, and likely produced increased sensitivity to environmental perturbations, then it is entirely expected that “inter-group competition” would emerge as one of the ways in which the carrying capacity would maintain a “check” on numbers. Sedentary agriculturalists must scramble for scarce resources. There’s no running off, at least at this stage of social complexity.

The fact that the LBK turned on each other should condition our understanding of how the transition to the Corded Ware may have occurred. The Y chromosomes of the LBK period are very different from what we find in Bronze Age Europe. The most reasonable model I believe is that these lineages did not go silently into the night. As they did to each other, so was done unto them. In J. R. R. Tolkien’s work there are allusions to the coming Fourth Age of Middle Earth, an age of men. The rise of agricultural mass society was the age of men in our world. Hunter-gatherer societies were no idyll, but due to their small scale, and complementarity in economic production, the relationship between the sexes was not one of male domination, where women were property to be traded as chattel. But concentrated and sedentary units of economic production that arose with village life became an inevitable target of extraction from collective groups of males, who translated their significant superior upper body strength into a reign of coercive terror. That coercion was translated into reproductive success, which is evident in the explosion of a finite set of Y chromosomal lineages on the order of ~5,000 years ago. The common R1a1a ancestor of Daniel MacArthur and myself was the original O G thug.

In evolutionary genetics R. A. Fisher introduced the idea that when selection pressures come to bear upon a population, large effect mutations may increase rapidly in frequency to increase population mean fitness. But, these mutations are not without cost, one reason that they were likely at low frequency in the first place. For example, one of the most well known adaptations to malaria famously has a very large segregation load in terms of a recessive disease. Evolutionary theory predicts over time that the adaptation will be less genetically disruptive. New mutations which allow for adaptation without the costs may emerge, or, other mutations may arise to “mask” and “modify” the deleterious effect of the initially favored allele.

When John Maynard Keynes purchased the papers of Isaac Newton he was shocked at the proportion of the great physicists writings devoted to matters occult and esoteric. Keynes declared that Newton was the ” last of the magicians, the last of the Babylonians and Sumerians, the last great mind which looked out on the visible and intellectual world with the same eyes as those who began to build our intellectual inheritance rather less than 10,000 years ago.” In opening the new age with his beautiful system of rational science, Newton nevertheless reflected an ancient ethos which persisted down into the modern period.

The Jewish people have been critical in the development of a universal ethical monotheism in the West, part of the broader evolution away from the supernatural systems of the Bronze Age that occurred across the Axial Age. But the Hebrew Bible preserves within it a world far removed from the divine Logos, a God of law and morality. The angry and jealous sky god of the Hebrews also enjoins upon them genocide of other tribes. Though the Hebrew Bible is pregnant with the possibilities of religious ethical universalism, the voice of the prophets’ righteous indignation raw with rage alive in our age, and channeled through the gentler voices of Hillel and Jesus, it also is a record of a parochial and peculiar people, who wash their hands of their atrocity by attributing it to the capricious and vindictive will of their god. If Moses and Joshua did exist, they almost certainly would have more in common with the war-chiefs of early Neolithic Europe, 4,000 years before their time, than men such as Constantine, who 1,300 years later promulgated a universal religion for a universal empire.

51w0iMybWyL._SY344_BO1,204,203,200_ Ancient Egypt, with its autocratic god-kings, was arguably one of the end-points of the Neolithic experiment with mass culture and ideology. So were Shang China and Mycenaean Greece, with their human sacrifices to propitiate the gods. Increasing primary productivity by an order of magnitude, which farming did, resulted in the emergence of huge amalgamations of humanity, and we as a species are culturally creative enough to have come up with adaptations. Literacy, cities, and social stratification, were all responses to the stresses and pressures that the opportunity of mass society presented. The emergence of powerful menacing and extortionate patrilineages was another. This was a world of gangs, thugs, and the question was not whether you would become a thug, it was whether you would be a thug or a victim of a thug. They were necessary, inevitable, cultural mutations against the background pressures that agricultural imposed upon humanity.

But as per Fisher’s model, mutants with deleterious consequences invite their own response. They are tamed and civilized by a scaffold of modifiers. The brutal gods which were but reflections of human vice and caprice were drafted in the service of primal human psychological impulses forged during the Paleolithic, reciprocity and egalitarianism arose against the background of brutality beyond imagining unleashed by the social dislocation that was a consequence of agricultural society. The men and women shaped by the Hebrew prophets and Christian Church Fathers, the rishis of the Upanishads and the Chinese sages, they are all closer to us 2,000 years later, then they were to their own forebears only a few hundred years earlier in their own past.

These models operate in the world between one of naive innate cognitive reflexes and pure cultural inventions generated without reference to the functional constraints of our minds and environments. The independent experiment of the Aztec Mesoamerican society suggests that the same stage of brutal social order that had occurred during the Neolithic was playing out in the New World. The Aztecs were engaging in ritual cannibalism and human sacrifice in a manner not seen in Old World civilizations since the Bronze Age. Some inventions are inevitable, emergent properties of the intersection of our biobehavioral toolkit and our species’ incredible cultural flexibility. Though we may believe ourselves to be far beyond the LBK people, the Nazi gas chambers or the more recent events in Rwanda suggest that the same mental reflexes of coalition-building and competition can be co-opted toward organized violent ends even today. Peace is possible, but violence is always imaginable.

Addendum: This Azar Gat article argues for the reality of war among hunter-gatherers, extensively citing what we know about Australian Aboriginal culture on the eve of European settlement. It would indicate that the only thing separating our Pleistocene ancestors from ourselves in terms of violence would be scale and organization, with ideology a novel handmaid.

 
• Category: History, Science • Tags: Indo-Europeans, War 
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51IQSePVDRL._SY344_BO1,204,203,200_ If there is one Peter Heather book you should read because it is timely, it is Empires and Barbarians: The Fall of Rome and the Birth of Europe. In it Heather makes an apologia for a revisionist view which resurrects some aspects of the old folk migration theories, and understandings of the arrival of barbarians into the collapsing Roman order of the middle of the first millennium. This is in contrast to the conventional view of modern archaeologists and historians which posits that the barbarian invasion was more a change of power to the elites, with the emergence of ethnic identities and coalitions almost in an ad hoc fashion among groups of mercenaries who took control from their paymasters. Heather does not posit total replacement of the indigenous population. In fact, it turns out in the best case scenario for such an event, in what became Anglo-Saxon England, the genetic data does not support such a proposition. Rather, there was an amalgamation between a culturally dominant intrusive minority, and the indigenous majority. The evidence from the rest of Western Europe is much more equivocal, suggesting that the demographic impact of the barbarians was minor (this not the case in Eastern Europe, where the Slavic migrations are associated with signals of strong genetic correlations between recently settled populations in the wake of German and Roman declines on the eastern frontier).

Heather’s position is really one of moderation or the Golden Mean. It is rather like those who do not take a pure hereditarian or environmentalist position in behavior genetics. In Empires and Barbarians he marshals evidence which points to the reality that the barbarian groups entered the Empire as self-conscious tribal-ethnic entities, with whole families on the move, and that they were not created de novo within the Empire. This is not to deny the reality of cultural shifts in identity, with Roman elites in Gaul taking to trousers and referring to themselves as “Franks,” and German tribal leaders attempting to accrue to themselves the glamour and respectability of Romanitas. But the fundamental identities which are combined were distinct, and organic, not recently constructed and inchoate.

51sBGrYcIEL._SY344_BO1,204,203,200_ The most recent work from ancient DNA, which I wrote about extensively this weekend, support Heather’s contentions broadly, if not specifically. The evidence from prehistory indicates far more demographic disruption than during the fall of the Roman Empire. That is, folk wanderings were much more significant in prehistory than in antiquity. That probably has to due social-demographic changes that occurred in the first millennium before the Common Era, as ruling elites became decoupled from the population they ruled in many ways, though often bound together by a religious ethos. The fusion of the conquered and conquerors was a process made much more feasible by the emergence of “meta-ethnic”, to use Peter Tuchin’s terminology, religious ideologies which transcended folk boundaries.

The reality of new facts means that we need to reinterpret aspects of archaeology and myth in terms of the dynamics which are reflected by our new understanding. One side aspect of my writings on these topics is that many Indians are not very happy with the newest results, because they validate threads of a frankly colonialist model of an Indo-Aryan invasion. The model is that a European-like population invaded the Indian subcontinent, imposed the caste system, and imparted many aspects of high culture upon the natives. Despite racial mixture between the indigenous and intrusive elements, the higher castes and peoples of the Northwest had more Aryan ancestry.

This model wasn’t totally a figment of the British colonialist imagination, and had much to do with the discovery of the connection between Sanskrit and other Indo-European languages, such as ancient Greek and Latin. Yes, the theory was utilized for political ends, but the model itself reflects some factual realities on the ground. For reasons which I will not elaborate in detail, because I don’t know that much about it, and don’t care either, a dominant strand of Indian nationalism has turned against the Indo-Aryan model of intrusion. In fact this school has “flipped the script,” asserting that Indo-Aryans, and therefore Indo-European languages, are indigenous to the subcontinent.

Linguistic sanity indicated that this was always unlikely. But the idea that most of the ancestry of South Asians dates to the last Ice Age, that is, the Indo-Aryan invasion was demographically marginal, was a defensible position until recently. For example, in 2006 you have a paper such as Polarity and Temporality of High-Resolution Y-Chromosome Distributions in India Identify Both Indigenous and Exogenous Expansions and Reveal Minor Genetic Influence of Central Asian Pastoralists. Additionally, as individuals have pointed out to me ad nauseam, archaeologists are skeptical of mass migration because of lack of remains.

R Graphics Output First, a paper from 2006 is wildly out of date for this field. The methods used have been superseded. Instead of using one locus, the Y chromosome, and focusing on microsatellites (which have upsides and downsides), researchers now look at the whole genome. Probably the best paper to read on the latest results is Genetic Evidence for Recent Population Mixture in India. The authors date the admixture between two different lineages in the subcontinent to “1,900 to 4,200 years ago.” One of the lineages is rather like West Asians and Europeans genetically (ANI). The other is most closely related to Andaman Islanders (ASI), but diverged from this group on the order of 20 thousand years ago. It can be accurately stated the the ASI are part of a broader constellation of East Eurasian populations which diverged from West and North Eurasians 40 thousand years ago (and from Basal Eurasians even earlier). In other words, modern South Asians are a compound of two highly diverged lineages, which explains their ambiguous position in relation to West and East Eurasian “reference” groups. And, this fact means teasing apart the ancestry is relatively easy, though time elapsed means that for all practical purposes South Asians are now a cluster of their own in any analysis of human population structure (that is, though they can be decomposed into parental elements, that’s not practical or useful in most cases).

Screenshot - 06152015 - 01:39:27 PM And, not surprisingly, the ratio of the ANI/ASI ancestry varies as a function of caste and region. This is the hallmark of relatively recent admixture, maintained by social barriers in caste. Indian caste groups (really jati) exhibit runs of homozogosity in their genome which are signs of a low effective population. In other words, not too much gene flow across groups. The direction of admixture is exactly as predicted by the old theories, as if an intrusive population arrived via the Northwest, and imposed itself upon the indigenous groups.

Now, let me address the major objections:

1) What about the mtDNA, which shows that most of South Asian ancestry is indigenous, and not West Eurasian.

Answer: The migration may have been male-mediated. All this means is that gene flow from the northwest occurred mostly through males. Over multiple iterations this can replace much of the whole genome, while leaving the single mtDNA lineage intact. This is exactly what happened in Argentina.

2) What about Y chromosomal results, which imply that R1a1a is indigenous to South Asia

Answer: Much of that work has utilized a small number of SNPs or microsatellites. The issues of phylogeography and dating for Y chromosomes are such that we really have had low clarity until whole genome analysis era, which is occurring now. The fact is that the Y chromosomal phylogeny of R1a1a the world over exhibits the hallmarks of massive population expansion ~5,000 years ago. No one knows where the R1 lineages are truly from in a deep time sense, as it seems likely they were low frequency variants before their explosion.

3) What about various aspects of mythology and archaeology which don’t posit an invasion/migration?

Answer: First, the archaeologists have a big record now of being wrong in Europe. Though many archaeologists (and historians who drew upon their scholarship) were cautious, some were not, and stridently argued that demographic replacements were not seen or evident in the archaeological record. The ancient DNA has basically proven them wrong. So either their methods miss migration, or, their results are labile when ideology is applied. Second, in regards to the lack of Indo-Aryan memory of migration, the Greeks also had no memory of migration. Both Greeks and Indians (Indo-Europeans) can not simultaneously be indigenous. Clearly oral memory has an expiration date, and may not be reliable.

Perhaps the archaeologists are totally right. I can’t evaluate their scholarship. I can evaluate the genetics, and it is getting progressively more persuasive. Scientific disciplines often have conflicts, though they are ultimately resolved (see the age of the earth controversy between geologists and biologists vs. physicists in the late 19th century). The recent track record of archaeologists has not been the best in my opinion when it comes to broader theoretical implications.

Finally, I want to state that I am skeptical, or not convinced, that most of the ANI ancestry in South Asia is from the Indo-Europeans. The ANI ancestry might be a composite, and perhaps the dominant language of the first ANI people was Dravidian. The closest populations to the ANI seem to be the people of the Caucasus, though this may be a function of changes in West Asian genetics. The Indo-Aryans may have been one of the later peoples to arrive.

Additionally, the Indo-Aryans were not blonde Europeans. Blonde Europeans seem to have evolved in Europe in situ over the past 4,000 years due to the admixture of diverse threads, indigenous and exogenous. There are elements of European ancestry (particular EEF) which do not seem to be evident in West Asia or South Asia, suggesting that the origination of Indo-European was complex, and perhaps multi-faceted. The Indo-Aryans had an affinity to the peoples of Europe, but it is a rather circuitous connection. We haven’t unpacked all the details.

Where does this leave us? There are many Indians who are like Creationists, for whom the scholarship is simply a way to buttress their ideology. They may cite the scholarship, but it is to a great extent just a matter of dressing up their nationalistic preferences. For these the new results are not only threatening, they are irrelevant. They will continue to cite papers from ~2005, because at that time period the results were more congenial to them. There are others who will “update” their views. There is much updating and revision that will occur over the next few years, and ancient DNA may change our perceptions about South Asia again. So be it.

Addendum: Wendy Doniger’s The Hindus is a controversial book. Religious fanatics and nationalists have waged a campaign against it. It is a testament of their milder nature that Doniger wrote under her real name, something that might not have happened if she was writing about Islam. I read the book when it came out, and it wasn’t my cup of tea, but I didn’t see what the big deal was (of course, why would I? I’m an irreligious atheist). But today I now think back to the passages which refer to the contact between Aryans and non-Aryans. Within these folktales and mythological memories may actually be a record of interaction which we do not in general have from Europe of “first contact” between startlingly disparate worlds.

 
• Category: Science • Tags: Aryan invasion theory, Indo-Europeans 
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61LXo6U7a4L._SY344_BO1,204,203,200_ Reading Strange Parallels, Southeast Asia in a Global Context, I have begun to think about the differences between the eruption of Inner Asian nomads in the early modern period, and in prehistory. The author points out that the arrival of Mughals, and even to a greater extent the Manchu, to the ancient and dense civilizations of South and East Asia did not change the cultural substrate in the main. Yes, Turco-Persian Islamic (“Islamicate”) culture became both prestigious and relatively popular in South Asia. But it was, and still is, a minority tradition set against the indigenous religious system, bracketed under the term Hindu today. In Ching China the Manchu had an even less obvious effect. Arguably they assimilated to the Neo-Confucian mores of the Han elite far more than the Mughals did in India in relation to indigenous South Asian gentry.

The dynamics in this context always need to take into account the numbers of the conquerors in relation to the conquered. The Manchus were less than 1 percent of the population of their domain. The foreign Muslim elites (Turk, Afghan, and Persian, with some Arabs in South India) and their scions were never more than a few percent, at most, of the population of South Asia. These alien elites rested atop an extractive system which predated them (in some cases by thousands of years). It was an institutional arrangement that was useful in terms of subsidizing their lifestyles. China and India were attractive to the nomadic populations beyond the frontier because they were rich with people, and therefore resources which could be deployed in consumption as well as marshaled for war (the Mughals milked India to finance wars in Afghanistan).

There are other cases which are similar in terms of numbers. Both the Magyar and Bulgar incursions into Europe seem to have resulted in an Inner Asian elite acclimating itself on top of a broad mass of peasants, from which it extracted rents. Though the Magyars imparted their name and language upon the populace of Roman Pannonia, genetically their impact has been fairly marginal, if detectable. The Bulgars, who exist only as they contributed to the appellation Bulgaria and Bulgarians, lost their language, and to my knowledge their genetic impact was even fainter.

But there are other cases. Both the Turks and Arabs seem to have more substantial genetic contributions, even if on the peripheries it was very marginal. Vast eras of Central Asia once inhabited by Persian and related Indo-European peoples has become a hybrid zone of sorts between West and East Eurasian peoples thanks to the Turkic migrations. Differences between Muslim and non-Muslim populations in the Fertile Crescent are evident.

Which brings me back to the Indo-Europeans. Even if they were not nomads of a classical sort which emerged later on in history, they seem to have been agro-pastoralists. There is now circumstantial evidence for their impact all across Europe, especially the north and east. There is also likely evidence for substantial Indo-European admixture in India. Herodotus reported 2,500 years ago that India was the most inhabited land on the face of the earth. But was it so 4,000 years ago, during the later stages of the Indus Valley civilization?

I will admit I was not primed to accept the idea of mass replacement of indigenous populations in what would later become the Ecumene by populations form the steppe because of the later record of conquest, which was more a matter of elite replacement, than social turnover. But if the genetic data is correct we need to update our models. If the first farmers of Europe were marginalized by invading Indo-Europeans, could not the same have happened to some extent to the agriculturalists of South Asia, who descended from the people of Mehrgarh? The tension between the interior and littoral in Eurasia is an old one, but it seems to have evolved over time, from one of inter-group competition and meta-population dynamics (read: extinction), to exploitation by Inner Asian steppes of the human resources of the littoral. Social complexity and institutional robustness were the best long term investments for farmer populations against nomads, who always outmatched them in individual skill, and often in terms of the tanks of the ancient world, horses.

 
• Category: History • Tags: Indo-Europeans 
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Fst between ancient European populations
Ba_MN Be_LN Ben_LN Cor_LN EHG LBK_EN Mo_HG Sp_EN Sp_MN Sw_NHG Un_EBA WHG
Baalberge_MN
Bell_Beaker_LN 0.020
BenzigeroH_LN 0.021 0.004
CordedW_LN 0.032 0.008 0.007
EHG 0.082 0.041 0.038 0.034
LBK_EN 0.009 0.021 0.025 0.035 0.084
Motala_HG 0.083 0.057 0.053 0.060 0.048 0.093
Spain_EN 0.021 0.028 0.035 0.044 0.092 0.015 0.099
Spain_MN 0.014 0.022 0.026 0.035 0.077 0.016 0.081 0.009
SwedSk_NHG 0.066 0.050 0.048 0.054 0.058 0.078 0.034 0.084 0.074
Unetice_EBA 0.016 0.002 0.003 0.005 0.034 0.022 0.051 0.030 0.023 0.043
WHG 0.073 0.055 0.056 0.070 0.078 0.091 0.053 0.092 0.070 0.050 0.057
Yamnaya 0.054 0.016 0.013 0.011 0.028 0.052 0.063 0.060 0.053 0.062 0.012 0.076

It is often said that the meeting of Europeans and Amerindians in the 15th century is our best taste of what it would be like to meet aliens. The analogy is rather straightforward, as Amerindians were not part of the broad interactions between societies over the Holocene, as they had removed themselves from the scene of action ~15,000 years ago. The interaction resulted in conflict and synthesis. A new people in Latin America arose who were biologically, and to a lesser extent culturally, a compound between two very distinct antecedents. In North America the dynamic was more of replacement and marginalization, often due to organized collective action on the part of white settlers and their political systems.

41ZlHlsdYXL._SY344_BO1,204,203,200_ Above I have again placed a selection of Fst statistics from the preprint Massive migration from the steppe is a source for Indo-European languages in Europe. One must be careful about not over-interpreting these statistics. But, they give a good sense of genetic divergence between two populations, on average (I have uploaded all Fst values in an Excel file). In particular above I’ve focused on the ancient populations in the data set. EN is Early Neolithic, while MN is Middle, and LN is Late. HG is Hunter-Gatherer. I’ve bolded Fst values > 0.05, because to me that’s a pretty high threshold for serious genetic divergence. The LBK brought agriculture to vast swaths of Central Europe. These were the people who encountered the hunter-gatherers who had occupied the continent since the end of the last Ice Age. The genetic distance between these two groups is incredibly high. Between the generic “Western Hunter-Gatherer” population and the LBK the Fst value is 0.091. What does that mean? The Mala are a Dalit (outside the caste system) group from Southern India. The Fst between this group and WHG is 0.11. In other words, not that much greater than between WHG and LBK. The Fst between the Mala and the LBK, two populations separated by a continent and 7,000 years is 0.077, less than between WHG and LBK. The genetic distance between the Mala and the French is 0.051. Considerably less than between the WHG and LBK. The get a sense of how small the values between modern European groups are, the genetic distance between Lithuanians and Sardinians is 0.021. The genetic distance between Belarussians and the English is 0.004, or about 4% of the distance between WHG and LBK. My claim that the contact between the first farmers and the hunter-gatherers of early Europe was one of intercontinental scale interaction is justified by the fact that the genetic distance between Han Chinese and Europeans is about ~0.10, only marginally greater than between WHG and LBK.

These sorts of genetic distances on the ancient European landscape could only be due to massive cultural revolutions which produced demographic shifts which can in no way be modeled as continuous diffusion processes. The Bantu expansion is probably a good analogy for what happened in ancient Europe, it took a little over 1,000 years to traverse the whole continent of Africa, a much larger geographic zone than Europe. But a second issue that we must also focus on are the genetic distances between European hunter-gatherers. One could chalk up 0.078 between EHG and WHG to the fact that EHG has admixture from “Ancestral North Eurasians” (ANE) at a clip of ~40 percent. But even the distance between the Motala hunter-gatherers, who are only 15% ANE, and WHG is 0.053. The uniparental markers are strongly suggestive of a relatively small group expanding to fill Europe after the last Ice Age, they are overwhelmingly haplogroup I for Y chromosomes, and haplogroup U5 for females. So what gives with the high Fst values? This is far higher than any modern intra-European distances. One hypothesis that I think might be viable is that small effective population cranked up the genetic drift in these groups, whose marriage networks were sharply delimited by cultural fractionation as well as ecological constraints which reduced population density outside of narrow specially favored areas (e.g., marine environments). One consequence of excess drift is elevated Fst values, beyond what you might think would be plausible. I’m not specialist in cultural evolution, but my intuition tells me that hunter-gatherer groups can engage in fission rather rapidly, and these divisions may have enforced greater barriers to gene flow to relatively recently diverged groups than is the norm in modern agricultural and post-agricultural societies.

Another issue that comes to mind are future analyses of ancestry tracts and linkage disequilibrium in these populations. The reason I bring this up is the fact that we need to distinguish between genetic differences due to standard workaday isolation by distance, and those produced by pulse admixture events (see Gideon Bradburd’s preprint for more elucidation of this issue). For various reasons outlined in the preprint above I’m convinced that a group like EHG, and later the hunter-gatherers of Scandinavia, did undergo admixture with a very different population during the early Holocene, after their expansion from the Ice Age refugia. But ultimately looking at patterns of this ancestry in the chromosomes as well as estimating a time since admixture would nail the coffin on this likelihood.

Finally, there’s the issue of “Nazi optics.” When word of these results started to percolate last spring a friend who is a prominent human geneticist blurted out “it sounds like the Nazis were right!” By this, he meant that this story of migrations, and demographic turnover, would be much more in place in Europe before World War 2. But there’s an immediate refutation of any attempt to Nazify these results: the Fst statistics made it clear that long term result of the clash of peoples in the early to mid Holocene has been amalgamation. The Bronze Age people of Europe, who gave rise to the historic nations, are the heirs of both the hunter-gatherers long indigenous to the continent, and disparate aliens, who arrived as strangers thousands of years ago.

 
• Category: History, Science • Tags: Indo-Europeans 
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Fst between selected populations
Armenian Bas Cord Czech EHG Eng Fren Gre LBK Lez Lith Sard Sin WHG
Basque 0.017
Corded_Ware_LN 0.023 0.025
Czech 0.011 0.009 0.015
EHG 0.067 0.060 0.034 0.043
English 0.011 0.008 0.014 0.002 0.045
French 0.009 0.006 0.015 0.001 0.048 0.001
Greek 0.004 0.010 0.019 0.004 0.057 0.005 0.003
LBK_EN 0.023 0.024 0.035 0.024 0.084 0.024 0.020 0.016
Lezgin 0.005 0.018 0.017 0.010 0.052 0.011 0.009 0.007 0.032
Lithuanian 0.019 0.014 0.016 0.003 0.043 0.006 0.006 0.010 0.034 0.015
Sardinian 0.014 0.013 0.033 0.013 0.074 0.012 0.009 0.009 0.015 0.021 0.021
Sindhi 0.016 0.032 0.029 0.022 0.057 0.023 0.022 0.020 0.049 0.014 0.028 0.035
WHG 0.086 0.062 0.070 0.056 0.078 0.058 0.058 0.070 0.091 0.082 0.053 0.074 0.087
Yamnaya 0.030 0.034 0.011 0.020 0.028 0.021 0.022 0.026 0.052 0.019 0.022 0.044 0.028 0.076

Update: Nick Patterson writes: Much ancient DNA genotype data is “pseudo-diploid” with just one allele given for a sample at each SNP. If you want to compute F_st values for such data, the easiest way is to run smartpca (see http://genetics.med.harvard.edu/reich/Reich_Lab/Software.html) with inbreed: YES set in the parameter file.

The downside is that this option requires at least 2 samples for each population while the default option (inbreed: NO) works on a single sample.


The above table is a selection of F ST values I culled from the preprint Massive migration from the steppe is a source for Indo-European languages in Europe. To get an intuition, the F ST value comparing Northern Europeans and Nigerians is ~0.15 using SNP data. Before I get to the meat of the argument above, let’s take that in for a moment. You see above that the pairwise values between WHG, “Western Hunter-Gatherers,” and the LBK, the first farmer culture of Central Europe, is on the order of ~0.10. That’s about the value between Europeans and East Asians today. We also know that it is possible there was a difference in color between the hunter-gatherers and the first farmers. The meeting of farmer and hunter-gatherer in early Neolithic Europe, and down to the Bronze Age, may best be thought of as analogous to a long term racial conflict and coexistence. Rather than a gradual wave of advance I envisage that the farmers hopped from point to point along fertile stretches of maritime littoral, and pushed their way up into the heart of Europe’s ancient forests by felling the wilderness around the great rivers which issue forth from the uplands. In a world of “isolation by distance” and “clines” this sort of recourse to a term like “race” would be anachronistic, but the model of genetic disruption being reported in these results using ancient DNA strong suggests punctuated demographic transitions across a wide range of localities which would result in biologically and culturally distinct groups persisting for many generations cheek by jowl. Over time admixture resulted in amalgamation, but it was almost certainly a millennia long process.

More specifically, the authors report unequivocally that the arrival of cultures like the Corded Ware in Northern Europe 4,500 to 5,000 years ago was accompanied by massive demographic replacement. Not only were these bands of warriors traversing the landscape, but it was a whole people on the move, men, women, and children. These were akin to the Goths fording the Danube and bursting into a new landscape of conquest. But the lands of the first farmers were not like those of Rome, heavily settled, further human cattle for the steppe agro-pastoralists to extract rents from. The preprint is not clear as to the timescale of the arrival of the eastern genetic influence across Southern Europe, but in the North the conclusion is without nuance or qualification: during the early years of the Egyptian Old Kingdom the lands of the north were being roiled by migration.

admix But in this post I want to turn the focus away from Europe for a bit. In the text they note that “An interesting pattern occurs at K=8, with all the late LN/BA groups from central Europe and the Yamnaya having some of the “light green” component that is lacking in earlier European farmers and hunter-gatherers; this component is found at high frequencies in South Asian populations….” I’ve edited and uploaded a version of the admixture plot. One must be cautious when interpreting these plots, but with all the other information in the paper it is quite informative. As far back as Noah Rosenberg’s 2005 paper, and later on in the blogger Dienekes Pontikos’ analyses, there were suggestions of affinities between a subset of Europeans and South Asians, as well as a connection to the Caucasus. In the F ST results above I note that the Sindhi population is closer to the Lezgins (Northeast Caucasians) and Yamna samples than they are to Armenians. The details are difficult to parse though. Otherwise they would have done so in this monumental paper.

I do want to add one final thing though. It’s been assumed in the past, including by me, that once farmers were established in a locale that future demographic perturbations were unlikely. By this, I mean that farmers are more useful alive to generate economic surplus for incoming elites than they are eliminated. But that is predicated on the idea of a complex specialized society where the elites view all non-elites in an almost Marxian sense of being objects of exploitation. It could very well be that this sort of cosmopolitan globalism only became common in complex societies later, and that in pre-state tribal groups even dense populations did not prevent extermination, because expropriation of vital resources necessary for survival was far more viable an option for these societies than exploitation.

 
• Category: History • Tags: Indo-Europeans 
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horsewheellanguage
Update: The preprint is out. End update

… and also after that, when the sons of God came in unto the daughters of men, and they bare children to them, the same became mighty men which were of old, men of renown.

Genesis, 6:4

A Battle-Axe

A Battle-Axe

An emanation from the one most high…uh, I mean, David Reich, has given his talk at Oxford. Thanks to Jean Manco we have a pretty good report of what he said. The core element seems to be that a paper will soon be published using ancient DNA results to conclude that Indo-European languages came to Europe from the Yamna culture of the Pontic Steppe ~4,000 years ago. Roughly, the argument laid out by David Anthony in The Horse, the Wheel, and Language: How Bronze-Age Riders from the Eurasian Steppes Shaped the Modern World. Where Reich’s conclusions differ from those that Anthony presented in the book is that this eruption from the Eurasian heartland resulted in a genetic-demographic transformation of Europe ~4,000 years ago. Manco confirms that the genetic data from the ancient sites that Reich’s team has access too indicate that the two dominant Y chromosomal haplogroups in Europe, R1b, and R1a, arrived with the Yamna people. This is not surprising, as previous ancient DNA exhibited a surprising death of these two lineages from both hunter-gatherer and early to late Neolithic populations in Europe. And, recent whole genome sequencing of Y chromosomes indicates that both R1a and R1b lineages across Eurasia have undergone recent explosive demographic expansion on a Holocene timescale (closer to 5 than 10 thousand years). In terms of total genome ancestry it looks as if the transition to Yamna successor cultures in Central Europe (e.g., Corded Ware) was accompanied by substantial genetic turnover. In the initial Corded Ware burial grounds 60-80 percent of the ancestry seems to have derived from the Yamna. The modern Yamna-derived proportion seems to be closer to ~50 percent in a region like Germany.

The Yamna themselves are a compound population, a mix of ancient hunter-gatherer groups (analogous to the modern Karelians), and an intrusive population with Near Eastern affinities, likely from the Caucasus. I am not clear whether the Near Eastern group had “Ancestral North Eurasian” ancestry, but the hunter-gatherers almost certainly did. The 1 to 18 percent “Ancestral North Eurasian” ancestry across continental Western and Central Europe today dates to the arrival of these Indo-European speakers. The very low fractions in Southern Europe, and its near absence in Sardinia, may suggest that the Indo-Europeans were demographically more significant in Northern Europe, even though they were clearly culturally effectual along the northern rim of the Mediterranean, and into Anatolia. In line with the Mal’ta results Manco relays that Reich believes that the R lineages, which are the dominant ones across Indo-European speaking populations from the North Sea down to South Asia, came into Europe via the Yamna, but ultimately derive from an “Ancestral North Eurasian” group.

220px-Yamna-en.svgThere are some phenotypic tidbits in the talk apparently. The Yamna were tall in terms of their genomic potential. Additionally, the very high frequency of lactase persistence may date to their arrival in Europe (there is some lack of clarity here). I doubt the high lactase persistence frequency and genotypes which result in greater final height are together by coincidence. Large people need a larger nutrient pipe, and adult digestion of lactose sugar would enable that.

There are two aspects which are not widely address in this talk. First, what was the exact dynamic of how the Indo-Europeans replaced the original populations? The idea of “demic diffusion” by waves of “demographic advance” promoted by Colin Renfrew seem to gradual and continuous to be responsible for this. This is basically an argument predicated on individual fitness, summed over groups. In this case I suspect that a better analogy may be the future that Genghis Khan had in mind for Northern China before his adviser Yelü Chucai dissuaded him: the North European plain was cleared out of people and turned over to pastureland. Genghis Khan and his Mongols were convinced of the value of Chinese as tax paying peasants, who could support the Mongol elite with their surplus. I suspect in a pre-state society such considerations were less relevant, as the institutional frameworks which would allow for the smooth absorption of subordinate groups were less elaborated, or even non-existent.

Hittites

Hittites

In both the Late Neolithic and after the Bronze Age the Reich group alludes to a return of the primal populations which were marginalized by the farmers, and later the Indo-European agro-pastoralists. One way to look at this is that there were larger migrations which were overlain upon the palimpsest. But, I believe one might also consider a model whereby there is ascertainment bias in the sorts of burial sites being explored and sampled, and one might be witnessing a patchy occupation of the landscape by intrusive cultures. For example, the newcomers might monopolize the rich bottom-lands for thousands of years, but huge swaths of the hinterland might be occupied by marginalized and less developed people, who over time drift into the core and become culturally absorbed. Instead of imagining the expansion of these people as purely ones of a vast uniform wave front, it might be better to conceptualize them as penetrating into virgin territory along the optimal avenues of settlement, and producing a patchy archipelago of habitation.

Second, there is the issue that though Reich and company focus on Indo-Europeans and the Yamna culture, the genetics leaves may loose threads that are difficult to tie back up. At ASHG Mait Metspalu express to me some misgivings about the term “Ancestral North Eurasian.” How do we truly know the locus and distribution of this ancestral component across Eurasia ~10,000 years ago? The Kalash of Pakistan exhibit signals of admixture with this group as high as Northern Europeans, so it is not limited to West Eurasia proper. The highest fractions today seem to be found in the North Caucasus, among many groups which are not Indo-European. If R1a was brought by Indo-Europeans to Europe, it is harder to conclude that this was the case in South Asia. Though the frequency of these lineages is higher in the Indo-Aryan North, there are relatively high fractions of R1a even among some South Indian tribal populations. R1b is found in appreciable fractions in Sardinia and among the Basques (one argument for the old idea that R1b was the legacy of European hunter-gatherers!). Obviously some of this could be due to admixture between Indo-Europeans and non-Indo-Europeans. But I think a major issue here is that Indo-European groups were a synthetic population which arose in a world where there were many synthetic populations, with ancient and recent affinities to them. I doubt the “Ancestral North Eurasian” ancestral component was limited purely to Indo-Europeans. So it seems unlikely that the R1 lineages would be purely Indo-European, even if recent expansion of some of their sub-lineages is a function of the Indo-European cultural explosion.

Of course there are only so many ancient DNA samples one can retrieve from a finite number of sites. The age of new genomic discoveries will start to close over the next few years as the paleo-demography inferred will start to exhibit some predictable solidity. That means that a deep knowledge of the archaeology, and what history there is, is essential.

 
• Category: History, Science • Tags: Corded Ware, Indo-Europeans 
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horsewheellanguage David Reich and Nick Patterson come down in favor of the steppe as the ur-heimat of the Indo-Europeans, at least those who migrated into Europe, in a recent abstract:

We generated genome-wide data from 65 Europeans who lived between 8,000-3,000 years ago by enriching ancient DNA libraries for a target set of about 390,000 single nucleotide polymorphisms. This strategy decreases the sequencing required to obtain genome-wide data from ancient DNA samples by around 1000-fold, allowing us to study an order of magnitude more individuals than previous studies and to obtain new insights about the past. We show that in western Europe, the farmers of both Germany and Spain >7,000 years ago were descended from a common ancestral stock. These farmers did not replace the earlier hunter-gatherers, but continued to mix with them, leading to a resurgence of hunter-gatherer ancestry in both Germany and Spain ~1,000-2,000 years later. In eastern Europe, the hunter-gatherers of Russia >7,000 years ago were distinct from those of the west, having an increased affinity to a ~24,000 year old individual from Siberia, but this affinity was reduced by ~5,000 years ago in the Yamnaya steppe pastoralists because of admixture with a population of Near Eastern ancestry. Western and Eastern Europe collided ~4,500 years ago with the appearance of the Corded Ware people in Central Europe, who derived at least two thirds of their ancestry from an eastern population closely related to the Yamnaya. The evidence for mass migration into Europe thousands of years after the arrival of agriculture, in combination with linguistic and archaeological data, makes a compelling case for the steppe as a proximate source for the spread of Indo-European languages into Europe.

This is broadly the same data which Iosif Lazaridis presented at ASHG 2014. So this itself is not new. But what I would like to draw your attention to are two posts over at Eurogenes, Ancient DNA points to the Eurasian steppe as a proximate source for Indo-European migrations into Europe, and Yamnaya genomes are a 50/50 mix of eastern Euro foragers and something else ANE-rich. Nick Patterson actually weighed in over in the comment thread for the first post. A comment in the second post was especially amusing:

Over 400 comments on an abstract? You may need to start a forum when the actual paper is released, David.

insearchof Yes, there were over 400 comments on the first post. It shows you how passionate people get about this issue. Some of the associations within this field are of a racialist nature. The Journal of Indo-European Studies was founded by Roger Pearson, though today it is edited by the respectable J. P. Mallory. This is not to say that all of those enthusiastic about this topic are quite so “out there,” but it’s quite emotional.

Until the paper itself comes out I suggest readers bone up on the archaeology, because there’s a wealth of that out there already. From what I recall the Samara samples were form David Anthony, and his The Horse, the Wheel, and Language: How Bronze-Age Riders from the Eurasian Steppes Shaped the Modern World is basically required reading in my opinion if you are interested in this issue. Also, Mallory’s older In Search of Indo-Europeans is probably worth reading as well. We live in interesting times indeed!

 
• Category: History, Science • Tags: Indo-Europeans 
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IE-speaking West Europeans are West Asian-admixed relative to Non-IE speaking Basques. Dienekes explicitly confirms what seems obvious using ADMIXTURE. When I get a chance I’m going to see if this difference is evident when comparing some South Indian (non-Brahmin samples) I have against Gujaratis. For what it’s worth I am told that ADMIXTOOLS will be out this week.

(Republished from Discover/GNXP by permission of author or representative)
 
• Category: Science • Tags: Genetics, Indo-Europeans 
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In response to my post below a friend emailed me the above sentence. As I suggest below it sounds crazy, and I don’t know if I believe it. But here’s an abstract from the Reich lab from June:

Estimating a date of mixture of ancestral South Asian populations

Linguistic and genetic studies have demonstrated that almost all groups in South Asia today descend from a mixture of two highly divergent populations: Ancestral North Indians (ANI) related to Central Asians, Middle Easterners and Europeans, and Ancestral South Indians (ASI) not related to any populations outside the Indian subcontinent. ANI and ASI have been estimated to have diverged from a common ancestor as much as 60,000 years ago, but the date of the ANI-ASI mixture is unknown. Here we analyze data from about 60 South Asian groups to estimate that major ANI-ASI mixture occurred 1,200-4,000 years ago. Some mixture may also be older—beyond the time we can query using admixture linkage disequilibrium—since it is universal throughout the subcontinent: present in every group speaking Indo-European or Dravidian languages, in all caste levels, and in primitive tribes. After the ANI-ASI mixture that occurred within the last four thousand years, a cultural shift led to widespread endogamy, decreasing the rate of additional mixture.


The most likely candidate population for an admixture event in the Indian subcontinent within such a time frame are Indo-Aryans. But, it does make some sense in light of the fact that the Northwest Eurasian variant of the lactase persistence allele is found in India, as that is presumably a relatively new variant. Let’s assume that in fact Indo-Aryans did arrive in India within this time frame, and were demographically numerous enough that they left a population genetic stamp. What is the probability that they did not do the same in Europe? I’d say that’s low. In other words, if the above results are correct, that Indo-Aryans had a significant impact on already heavily populated South Asia, then it stands to reason that the same would be true for Europe. Why hasn’t this signal been as easily detected? I Assume it’s because the Indo-Europeans were genetically closer to non-Indo-European Europeans in the first place.

This sort of phenomenon may explain the recent divergence between Tibetans and Chinese. Linguistically the two populations are very different, and it is hard to credit that Tibetan and Chinese dialects diverged within the last ~3,000 years. But it makes much more sense if the Han demographic radiation was from a set of genetically similar, but culturally diverse, populations. Though gene flow maintained a degree of coherency, it may be that there were deep linguistic fissures across the North China plain before the rise of the Han. Similarly, the peoples of the Caucasus exhibit a lot of linguistic diversity, far more than you’d predict from simple genetics. The Indo-Europeans in Europe may have assimilated linguistically very different people, who were genetically quite similar. In India, they may have assimilated linguistically very different people, who were also genetically distinct.

Addendum: I do believe that the highest probability is that Europe and India saw multiple intrusive populations after the rise of agriculture. So the larger proportion of the ANI signal probably derives back to the early West Asian farmers.

Image credit: Wikipedia

(Republished from Discover/GNXP by permission of author or representative)
 
• Category: History, Science • Tags: Anthropology, Indo-Europeans 
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Dienekes P. is often rather laconic in commentary on the papers he links to, but of late he has “come out of his shell.” He has two posts which are important “weekend reading”:

- Population strata in the West Siberian plain (Baraba forest steppe)

- Hints of East/Central Asian admixture in Northern Europe

I freely admit that much of the conjecture here is above my pay-grade in terms of evaluation. But I do think it’s important think through. My “gut” tends to lean toward a “revenge of the Mesolithic” scenario promoted by some of Dienekes’ critics, but I don’t have a strong position.

(Republished from Discover/GNXP by permission of author or representative)
 
• Category: History, Science • Tags: Indo-Europeans 
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I don’t know the answer to the question posted in title above, and I’m moderately skeptical that he has. But I wanted to give him full credit in the public record if researchers confirm his findings in the next few years. You can read the full post at his weblog, but basically he found that a West Asian modal element in a north British (Orkney) and Lithuanian individual seems to be negatively correlated with a Northwest European modal element and positively correlated with Near Eastern and South Asian components on a genomic level across different models in ADMIXTURE (e.g., does “South Asian” at K = 5 tend to match “West Asian” at K = 8).

Two major concerns:


- I don’t have a good intuition for this method. Could this be an artifact of the algorithm?

- When you have a hypothesis in mind you can unconsciously seek out confirmatory points. As you can see in the comments below Dienekes and his interlocutors have given this issue much thought. Frankly, I found it difficult to follow a lot of the dialogue, and I follow this topic more than most.

It seems that at this point someone should do follow up analyses with other populations, assuming that the method is informative.

(Republished from Discover/GNXP by permission of author or representative)
 
• Category: History, Science • Tags: Anthropology, Indo-Europeans, Linguistics 
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Dienekes has a post up, The Bronze Age Indo-European invasion of Europe. The crux of his argument is as such:

But there is another component present in modern Europe, the West_Asian which is conspicuous in its absence in all the ancient samples so far. This component reaches its highest occurrence in the highlands of West Asia, from Anatolia and the Caucasus all the way to the Indian subcontinent. It is well represented in modern Europeans, reaching its minima in the Iberian peninsula….

Thanks to the public release of genetic data Dienekes has developed his theories in part out of his own analyses of said data. Though I’ve run fewer analyses, with smaller data sets, some of the same patterns jump out at me. In particular, there is a component which is modal in northern West Asia (e.g., the trans-Caucasian region) which seems to drop mysteriously between the French generally and French Basques, and the Basque vs. non-Basque Spanish samples. There are also similar, though not necessarily easy to map across the two regions, disjunctions in South Asia between geographically close Indian groups.


Ultimately model-based clustering algorithms and PCA is going to get us only so far. Remember that the clusters generated from these methods don’t give us reality as such, but particular patterns which map back to reality. You can’t read from cluster A to population X without a non-genetic interpretative frame. Nevertheless, I do think within the next few years we may solve the “mystery” of the demographic origin of Indo-European languages and culture through genetics.

First, we have to posit a hypothesis in the fashion which Dienekes proposes. That is, Indo-European languages began to spread rapidly ~5,000 years ago from a small core population. This rapidity leads to both cultural integrity, and some genetic signal, which spans Indo-Eurpean groups. There are two dimensions, time and space. As Dienekes notes ancient DNA data points will get thicker. If the West Asian component in European ancestry begins to show up >5,000 years B.P., that’s going to be highly suspicious. Second, our data set of extant populations is going to get thicker. You’ll be able to independently contrast relatively similar Indo-European vs. non-Indo-European populations. For example, Sinhala vs. Tamil and Swede vs. Finn. By and large the two tips of the two clades should exhibit genetic similarity, but if the thesis of Indo-European demographic expansion is correct then you’re going to see a subset of matches which correlate with language family, and not proximity. You can then construct a “synthetic” genome from these matches across independent pairs. Finally, you can compare that genome to present populations and ancient DNA samples.

Addendum: I assume that methods like looking at identity-by-descent tracts are going to be important, even though recombination will have broken apart the regions quite a bit.

(Republished from Discover/GNXP by permission of author or representative)
 
• Category: Science • Tags: Anthropology, Indo-Europeans 
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As must be obvious, I think now that the spread of Indo-European languages had some demographic impact. It wasn’t analogous to the spread of English to Jamaica, or the existence of French as an official language in Congo-Brazzaville. Because of this, I now believe it is possible in the near future that scientists will reconstruct the genome of the original Indo-Europeans. How?

1) Find the intersection of genetic segments on the chromosomal level which share identity-by-descent between widely separated Indo-European groups. For example, Greeks, Swedes, and Punjabis.

2) Check to see which of these intersecting elements is not found in nearby non-Indo-European groups. For example, Basques, Finns, and non-Brahmin South Indian Dravidian speakers. At least to an appreciable frequency.

My current supposition is that proportionally this component won’t be preponderant in most places, but, it will be significant. By reconstructing an Indo-European genome we may actually have the ability to ascertain the population’s urheimat, as we can compare its genetic distance to extant populations.

(Republished from Discover/GNXP by permission of author or representative)
 
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The image above is adapted from the 2010 paper A Predominantly Neolithic Origin for European Paternal Lineages, and it shows the frequencies of Y chromosomal haplogroup R1b1b2 across Europe. As you can see as you approach the Atlantic the frequency converges upon ~100%. Interestingly the fraction of R1b1b2 is highest among populations such as the Basque and the Welsh. This was taken by some researchers in the late 1990s and early 2000s as evidence that the Welsh adopted a Celtic language, prior to which they spoke a dialect distantly related to Basque. Additionally, the assumption was that the Basques were the ur-Europeans. Descendants of the Paleolithic populations of the continent both biologically and culturally, so that the peculiar aspects of the Basque language were attributed by some to its ancient Stone Age origins.

As indicated by the title the above paper overturned such assumptions, and rather implied that the origin of R1b1b2 haplogroup was in the Near East, and associated with the expansion of Middle Eastern farmers from the eastern Mediterranean toward western Europe ~10,000 years ago. Instead of the high frequency of R1b1b2 being a confident peg for the dominance of Paleolithic rootedness of contemporary Europeans, as well as the spread of farming mostly though cultural diffusion, now it had become a lynch pin for the case that Europe had seen one, and perhaps more than one, demographic revolutions over the past 10,000 years.

This is made very evident in the results from ancient DNA, which are hard to superimpose upon a simplistic model of a two way admixture between a Paleolithic substrate and a Neolithic overlay. Rather, it may be that there were multiple pulses into a European cul-de-sac since the rise of agriculture from different starting points. We need to be careful of overly broad pronouncements at this point, because as they say this is a “developing” area. But, I want to go back to the western European fringe for a moment.


As I stated above the Basques were long used as a Paleolithic “reference” by historical geneticists. That is, the deviation of a population from the Basques would be a good measure of how much admixture there had been from post-Paleolithic sources. Connections between Iberian populations and those of western and northern Europe were used to trace expansions out of the ecological refuges of modern humans during the Last Glacial Maximum ~20,000 years ago. Just goes to show how reliant we are on axioms which are squishier than we’d like to think.

Last fall I posted a result from Dodecad on the difference between French and French Basques (both from the HGDP). I’ve replicated this myself a few times now too:

The striking aspect is that the Basque are less cosmopolitan than the other French. This is evident in most of the runs of the HGDP Basque; they just have a “simpler” genetic heritage than other Western Europeans. Today Dienekes posted some results from the IBS Spanish data set in the 1000 Genomes. He suggests there are clearly a few Spanish Basques in there (I’ve highlighted them):

Recall that the Basques were exempt from inspection for “cleanliness of blood”, because they were presumed to lack Jewish or Moorish ancestry by virtue of being Basque. It seems that the Spanish IBS sample, like the Behar et al. Spaniards and Portuguese, do have some Moorish genetic imprint. This is not too surprising. The Moriscos might have been expelled in the early 17th century, but not before the majority had converted to Christianity over the centuries (in fact, some of the most virulent anti-Morisco partisans had Moorish ancestry themselves, and were particularly tainted by association with the remaining culturally unassimilated crypto-Muslims). All that being said, I suspect that the “West Asian” ancestry amongst the majority of the Spaniards is not due mostly to the Arab period (when of the majority of the settlers probably were Berbers or Arabicized Berbers), but to population impacts prior to that. By the time of the Roman conquest much of Spain was Celtiberian. I have low confidence in this assertion, but I am coming to believe that the Indo-Europeans brought a mix of East European and West Asian ancestry (or at least those two distinct strands which tend to shake out of ADMIXTURE in a broad array of European samples) to western Europe.

On a related note, Wave-of-Advance Models of the Diffusion of the Y Chromosome Haplogroup R1b1b2 in Europe:

Whether or not the spread of agriculture in Europe was accompanied by movements of people is a long-standing question in archeology and anthropology, which has been frequently addressed with the help of population genetic data. Estimates on dates of expansion and geographic origins obtained from genetic data are however sensitive to the calibration of mutation rates and to the mathematical models used to perform inference. For instance, recent data on the Y chromosome haplogroup R1b1b2 (M269) have either suggested a Neolithic origin for European paternal lineages or a more ancient Paleolithic origin depending on the calibration of Y-STR mutation rates. Here we examine the date of expansion and the geographic origin of hgR1b1b2 considering two current estimates of mutation rates in a total of fourteen realistic wave-of-advance models. We report that a range expansion dating to the Paleolithic is unlikely to explain the observed geographical distribution of microsatellite diversity, and that whether the data is informative with respect to the spread of agriculture in Europe depends on the mutation rate assumption in a critical way.

Really I’m waiting for more ancient DNA. These sorts of studies are starting to feel like rewarming cold pizza. Edible, but suboptimal. Next, Phylogeography of a Land Snail Suggests Trans-Mediterranean Neolithic Transport:

Background
Fragmented distribution ranges of species with little active dispersal capacity raise the question about their place of origin and the processes and timing of either range fragmentation or dispersal. The peculiar distribution of the land snail Tudorella sulcata s. str. in Southern France, Sardinia and Algeria is such a challenging case.

Methodology
Statistical phylogeographic analyses with mitochondrial COI and nuclear hsp70 haplotypes were used to answer the questions of the species’ origin, sequence and timing of dispersal. The origin of the species was on Sardinia. Starting from there, a first expansion to Algeria and then to France took place. Abiotic and zoochorous dispersal could be excluded by considering the species’ life style, leaving only anthropogenic translocation as parsimonious explanation. The geographic expansion could be dated to approximately 8,000 years before present with a 95% confidence interval of 10,000 to 3,000 years before present.

Conclusions
This period coincides with the Neolithic expansion in the Western Mediterranean, suggesting a role of these settlers as vectors. Our findings thus propose that non-domesticated animals and plants may give hints on the direction and timing of early human expansion routes.

So basically the snail hitched a ride from Sardinia to Algeria to France. I don’t think this is that surprising. First, it seems pretty obvious that a lot of the cultural expansion in the prehistoric period did not consist of the fission of villages along a continuous wave of advance, but involved leap-frogging to suitable nuclei from which the populations expanded. Imagine a rising flood where the lowest zones are inundated first, and then the higher peaks. Additionally, we shouldn’t presume that these expansion events were without conflict and institutional support. Consider that the expansion of farming across much of southern European Russia and Ukraine could only occur after the state had pacified, expelled, or assimilated, the mobile Turkic populations which were wont to extract unsustainable rents out of isolated and vulnerable peasant populations.

Finally, what’s up with the strong north-south differentiation across the Mediterranean basin, peaking in the west? It’s as if there were two waves of demographic and cultural advance which laid the ground work, and later perturbations haven’t disrupted that bedrock. It suggests to me the critical importance of lateral coastal transport in connecting cultural colonies, as opposed to more long distance jumps across the open sea. The latter were probably important for the transport of luxury goods and the exchange of memes, but not so much for the exchange of genes.

(Republished from Discover/GNXP by permission of author or representative)
 
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The following passage is from the epilogue of The Real Eve: Modern Man’s Journey Out of Africa by Stephen Oppenheimer:

In this book I have offered a synthesis of genetic and other evidence. Everything points to a single southern exodus from Eritrea to the Yemen, and to all the non-African male and female gene lines having arisen from their respective single out-of-Africa founder lines in South Asian (or at least near the southern exit). I regard the genetic logic for this synthesis as a solid foundation, and I have based the rest of my reconstruction of the human diaspora upon it. Obviously, the ‘choice’ of starting point (mine or theirs) determined all the subsequent routes our ancestors and cousins took. Tracing the onward trails is only possible as a result of marked specificity in regional distribution of the genetic branches The geographic clarity of both male and female gene trees is a big departure from the fuzzy inter-regional picture shown by older genetic studies. The degree of segregation of lines into different countries and continents is in itself good evidence that once they got to their chosen new homes, the pioneers generally stayed put, at least until the Last Glacial maximum forced some of them to move. This conservative aspect of our genetic prehistory also provides a partial explanation for the fact that when we look at a person, we can usually tell, to the continent, where their immediate ancestors came from, and underlies differences that some of us still call ‘race.’

Oppenheimer wrote the above in the early aughts, as his book was published in 2003. Much of this is generally in line with the ‘orthodoxy’ of the day. I believe that Oppenheimer’s assertion that there was one southern migration out of Africa by anatomically modern humans has gained some advantage over the alternative model of two routes, northern and southern, over the past ten years (Spencer Wells’ The Journey of Man sketches out the two wave model). Other assertions and assumptions have not stood the test of time. In particular, I would contend that generally the ‘conservative aspect of our genetic prehistory’ can no longer be taken for granted. Specifically, it seems likely now that much occurred after the Ice Age and during the Neolithic.


420px-AGMA_HérodoteThe false inferences of the early aughts were due to two primary problems. First, they relied heavily on the powerful new techniques of extraction and analysis of uniparental ineages; the male and female direct line of descent. Concretely, mtDNA and the nonrecombintant portion of the Y chromosome. The lack of recombination allows for relatively easy reconstruction of phylogenies assuming a coalescent model. Second, the inferences attempt to make connections between the patterns of variation in modern populations, and what one may infer about the past from those patterns. Obviously constructing a phylogeny, or plotting haplogroup frequencies as a function of geography, is rather straightforward science. But using these results to generate inferences of the past is often more of an art than a science, and implicit assumptions lurk behind the causal chains. Consider for example the utilization of modern Anatolian (i.e., Turkish) genetic variation as a reference for the expansion into Europe of Neolithic farmers from the Near East. This of course presumes that modern Anatolians are a good proxy for ancient Anatolians. There are various suggestive reasons for why this is a plausible assumption, but assemble enough plausible assumptions, and rely on their joint likelihood, and you construct a very rickety machinery of possibility.

In early 2007 I began to have serious doubts about the orthodoxy of genetic conservatism. The primary trigger was the story of the Etruscans. Here is the crux of the issue: there are two models for the origins of the Etruscans, first, that they were the pre-Indo-European autochthons of Italy, or, that they were the migrants from the eastern Mediterranean, in particular Anatolia. The second may seem an outlandish hypothesis, but there were several tendrils of evidence to support it. But perhaps the ‘support’ which weighed most against it is that the fact that the Anatolian model has an ancient source, the Greek historian Herodotus. I should perhaps put historian in quotes as well, because Herodotus is often viewed more as a repeater of myths, and derided by some as the ‘father of lies’ (in this he stands in sharp contrast to contemporary perceptions of the ‘modern’ Thucydides, though revisionists have begun to challenge this narrative). In contrast, the model that Etruscans are indigenous to Italy, and that their ‘exotic’ foreign traits were simply acquired through trade and cultural diffusion, dovetailed well with the post-World War II ‘pots not peoples’ paradigm. That cultural change was ubiquitous, while at the same time populations were immobile. It was boring, prosaic, and conservative, and so an ideal null hypothesis.

But here it turns out that Herodotus was right, and archaeologists were wrong. Genetic analysis of modern Tuscans from isolated villages shows that some are surprisingly closely related to extant eastern Mediterranean lineages. Genetic analysis of Tuscan cattle showed that they were surprisingly closely related to extant eastern Mediterranean lineages of cattle. Finally, extraction of ancient Etruscan DNA showed that they were closely related to extant eastern Mediterranean lineages. The overlap was often with Anatolia, and combined with fragmentary linguistic and archaeological data, the evidence clearly points to an exogenous origin for the Etruscans. The boring null hypothesis was wrong. After these genetic stories gained prominence I went and reread recent archaeological texts on the Etruscans, and there were many models which showed exactly how Etruscan cultural uniqueness derived back to prehistoric Italy. It seems in hindsight that the prior assumption served as an interpretative filter, and people saw patterns that they were primed to see based on what they ‘knew’ to be the history of prehistoric and early Iron Age Tuscany.

Of course to refute the primacy of Oppenheimer’s conservative model of genetics one has to offer more examples than that of the Etruscans, and in particular, examples which are of greater scope and weight. I believe those examples exist. In the early aughts based on the mtDNA evidence the likelihood was that South Asian genetic variation is by and large a product of changes wrought upon the basic elements extant in the region around the end of the last Ice Age. The Y chromosomal data was more confused, though it did imply a closer relationship to groups in western Eurasia. But based on the mtDNA Oppenheimer posited a model whereby India was the mother of all non-Africans, that is, all non-African lineages derived from roots within the Indian subcontinent before the Last Glacial Maximum. This is at sharp variance with colonialist narratives of an Aryan invasion of the subcontinent, and the subjugation of the natives by quasi-European overlords, who are the ancestors of the moder upper castes. The charged ideological import of this model is transparently obvious.

Unfortunately the reality is likely more complex. I suspect that some form of Oppenheimer’s model is correct, insofar as South Asia was likely an important way station for modern humans as they left Africa, and pushed into other regions of Eurasia, on to Australasia and the New World. This interpretation does gain support from mtDNA, the direct maternal lineage. But a new analysis of South Asian genetic variation using a substantial proportion of the autosomal genome implies in fact that South Asians are possibly descendants of an ancient hybridization event between a native population with deep roots in the subcontinent, and a quasi-European population which was exogenous to the subcontinent.* Genetically the quasi-European population is quite close to northern Europeans, similar to the genetic distance between modern Finns and Italians, not trivial, but far closer than that between modern South Asians and Europeans. Was this the ancient Aryan invasion? I remain skeptical of this particular detail for various reasons, as I suspect that the history of the Indian subcontinent is in fact even more complex than has been assumed before (I think it is more likely that the quasi-Europeans came before the Indo-Aryans, who arrived late, and had a stronger cultural than genetic influence).

Finally, there is another region of the world where it seems likely that the old orthodoxies of genetic conservatism will be overthrown. That region is Europe. The scientific orthodoxy of deep time continuity is strong enough that it has percolated into the public consciousness, the leader of the British National Party even referred to the deep roots of white British in demarcating who he believed ‘indigenous people’ of the Isles were. But newer data is more supportive of the hypothesis that in fact Neolithic farmers who arrived from elsewhere are the likely ancestors of most Europeans, not the hunter-gatherers who remained after the Ice Age. Extraction of ancient DNA has yielded a set of results which simply are not explicable assuming the older models of genetic continuity, which were based on inferences made from modern population variation. If I had to hazard a guess, I would have some, though not high, confidence in the following story. First, the indigenous hunter-gatherers are assimilated or marginalized by waves of Neolithic farmers pushing out from the eastern Mediterranean. The demographic expansion does not necessarily sweep outward along a southeast-northwest axis, rather, it follows the Mediterranean and Atlantic fringes, as well as along river systems in the interior. Its impact is weakest in the northeast of Europe, where Middle Eastern crops are least suitable, and the natives have the most time to absorb the cultural toolkit of the newcomers so as to resist their advance. Second, and far later, there was another wave pushing out from the region of the Ukraine to the Volga, likely the ancestors of the Indo-Europeans. Tentatively I would contend that these were the carriers of the Kurgan culture, and also brought the allele for lactase persistence. Again, for ecological reasons the populations of the northeast Baltic and into the forests of northern Russia were most insulated from this push (and non-Indo-European languages persisted in Iberia down to Roman times, and specifically in the Basque-country down to modern times, though I suspect this is a function of distance). So modern European populations may be assumed to be tri-hybrid, first a synthesis of Middle Eastern farmers overlain upon the Paleolithic substrate, and second a synthesis of Indo-Europeans from the east overlain upon pre-Indo-European substrate. Unlike the case of India I suspect teasing out these patterns in modern populations is more difficult because the genetic distance between the three ancestral populations is far smaller than between the indigenous peoples of India before the quasi-Europeans arrived.

This leaves much of the world untouched by my speculations, but I believe showing that the genetically conservative null hypothesis is now in serious doubt in South Asia and Europe is sufficient to knock it from being a necessarily default assumption through which we must filter our interpretations. I do not believe that the reordering of human variation and the welter of population movement after the Ice Age was equivalent in effect to the Out of Africa migration, but I do believe that it was important enough to make the world of 2000 BCE very different from that of 15000 BCE in regards to genetic variation. In some cases, such as Central Asia from the Caspian to the Taklamakan the world of 2000 CE is fundamentally different from the world of 0 CE.

I will then end with a prediction, one in which I do not have much confidence, but which may no longer be wrong on the face of it with these new data in mind. Here is a passage from page 7 of Jared Diamond’s Guns, Germs, and Steel:

Initially, archaeologists considered the possibility that the colonization of Australia/New Guinea was achieved accidentally by just a few people swept to sea while fishing on a raft near an Indonesian island. In an extreme scenario the first settlers are pictured as having consisted of a single pregnant young woman carrying a male fetus…..

Let me stipulate that Diamond seems skeptical of the extreme model, but it illustrates the consensus that Australian Aboriginal populations are descended from the first settlers. That is, the modern populations of indigenous Australians are the direct descendants of those who swept Out of Africa along the fringe of the Indian ocean, through Southeast Asia, and arrived in Australia (more specifically, Sahul), on the order of 40 to 60 thousand years ago. From what genetic data I have seen this may be true. But I do not know of any extractions of ancient DNA, and it seems to me that the analysis of the phylogenetics of Australian Aboriginals is relatively sketchy. Therefore, I will suggest that within the last 10,000 years there has been a major new migration of people into Australia, and the modern range of genetic variation of Australian Aboriginals is significantly different from that of the populations of the Ice Age. I suggest this primarily because the dingo arrived within the last 10,000 years, more likely as recently as 4,000 years ago. With the expansion of the utility of ancient DNA extraction and analysis this question may be answered in the near future. I would still bet I’m wrong with the hypothesis I just offered, but I’m far less sure than I would have been 2 years ago.

Note: This post emerged from a conversation I had with Kevin Zelnio and Dave Munger.

* I say ‘quasi-European’ because the population may have origins outside of the boundaries of modern Europe at the Urals. Perhaps in western Siberia. Additionally, the idea of ‘Europe’ is relatively new, and exhibits little ancient cultural coherency.

Image source: Wikipedia

(Republished from Discover/GNXP by permission of author or representative)
 
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Razib Khan
About Razib Khan

"I have degrees in biology and biochemistry, a passion for genetics, history, and philosophy, and shrimp is my favorite food. If you want to know more, see the links at http://www.razib.com"