The population genomics of South Asia is complicated, and politics doesn’t make it easier

Many people have been sending me links to this article, By rewriting history, Hindu nationalists aim to assert their dominance over India. Here’s a key section:

The RSS asserts that ancestors of all people of Indian origin – including 172 million Muslims – were Hindu and that they must accept their common ancestry as part of Bharat Mata, or Mother India. Modi has been a member of the RSS since childhood. An official biography of Culture Minister Sharma says he too has been a “dedicated follower” of the RSS for many years.

Sharma told Reuters he expects the conclusions of the committee to find their way into school textbooks and academic research. The panel is referred to in government documents as the committee for “holistic study of origin and evolution of Indian culture since 12,000 years before present and its interface with other cultures of the world.”

Sharma said this “Hindu first” version of Indian history will be added to a school curriculum which has long taught that people from central Asia arrived in India much more recently, some 3,000 to 4,000 years ago, and transformed the population

There are several threads here. First, it is a fact that the ancestors of South Asia’s non-Hindus were Hindu. There are minor exceptions, such as the Parsis, who are ~75% Iranian. One can quibble as to whether many tribal and peasant populations were truly Hindu in a formal and explicit sense. But I think this is a semantic dodge. Muslims would recognize these beliefs and practices as Hindu, no matter if one was a Brahmin monk or a member of a tribe which still sacrificed animals.

I’ve looked at the genotypes of a fair amount of South Asians of Muslim background. The overwhelming (usually exclusive) proportion of their ancestry is South Asian. It’s a fact that the ancestors of non-Hindu South Asians were Hindu.

But, the article and a dominant theme in Hindu nationalism today are that distinctive historically important groups like Indo-Aryans are indigenous to South Asia. This is set against a narrative of invasions and migrations from the outside, which is presumed more friendly to a multicultural paradigm (I have a hard time keeping track of the political valence of all these things). To some extent, the reality of invasions and migrations cannot be denied, whether it be Alexander, the Kushans, or the various Muslim groups. But these historical invasions left little genetic imprint.

When 2009’s Reconstructing Indian Population History was published things changed for the impact of the earlier migrations. By the time the ancient Greeks were recording observations of India in Classical Antiquity, it was already noted as the most populous nation in the world. I was initially skeptical about the result in Reconstructing Indian Population History, that there was massive admixture between West Eurasians (ANI) and indigenous South Asians (ASI) because that would imply massive migration. Additionally, phenotypically the pigmentation genes didn’t seem to work out if the source population was European-like.

Nearly 10 years on we have a lot more clarity. Ancient DNA has changed our understanding of the past. Massive migrations were common. And, the pigmentation and genetic profile of modern Europeans is recent, within the last 4,000 years. The source population(s) for “Ancestral North Indians” (ANI) may not have been Europeans in the way we’d understand them. In fact, a follow-up paper, Genetic Evidence for Recent Population Mixture in India, hinted at two admixtures. There’s a fair amount of circumstantial evidence now that one component of “Ancestral North Indian” relates to West Asian populations and another component to the more classical steppe Indo-Aryans. The former is more widespread across the subcontinent than the latter, which is concentrated in the northwest and among upper castes.

I do understand Indians who want to interpret their own history through the lens of their own cultural priors. The problem is that genetic science has proceeded so fast in the last few years that many propositions which were speculative in the 20th century are testable in the 21st century. Some Hindu nationalist friends and acquaintances express embarrassment and worry about the track that Indian nationalists are going on. I don’t know what to say, but Americans have their own delusions and blithe acceptance of propaganda, so I’m not going to be one pointing fingers. Other Indians have told me via Facebook that they “believe in the results from the 2000s” (when they were more congenial to their viewpoints?). I guess that’s one strategy; just keep up with the science until it starts refuting your model.

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The Dravidianization of India

On this week’s The Insight Spencer Wells and I talk about the Indo-Aryan arrival to South Asia. This was recorded very early last summer, and I’m rather unguarded (it’s well before I had the piece published in India Today).

I think 2018 will finally be the year that a lot of South Asia will be “solved.” There has been some foot-dragging on papers and results, but that can only go so long.

All that being said I suppose I should make some suppositions I have arrived at on this topic more explicit, as in a discussion with an Indian friend he admitted had no idea about some of my views, though he reads this weblog when I expressed them. That’s because they are speculative and my confidence in them is weak, though you can infer my opinions if you look very closely.

The figure to the left is from Genomic insights into the origin of farming in the ancient Near East, a paper published about a year and a half ago. You see various South Asian populations being modeled as a mixture of four different source populations. The Onge are an Andaman Islander population (and the closest we can get to the aboriginal peoples of South Asia). Iran_N represents Neolithic Iranians, the canonical “eastern farmer” population. Steppe_EMBA represent Yamnaya pastoralists, who are themselves modeled as a mixture of Eastern European Hunter-Gatherers (EHG) and southern population which has affinities with the Iran_N cluster. EHG in their turn seems to exhibit ancestry from Western European Hunter-Gatherers (WHG), whose heritage dates to the late Pleistocene, and Ancient North Eurasians (ANE), who flourished in Siberia, and contributed ancestry to populations to the west and east (including the ancestors of Native Americans).

When I first saw this specific figure I was incredulous. I had long thought that “Ancient North Indians” (ANI) were a compound of two elements, one related to the farmers of West Asia (Iran_N), and the other steppe Indo-European (Steppe_EMBA/Yamnaya). But the fraction of Yamnaya/Indo-European/Indo-Aryan ancestry seemed far too high.

A few years later I am less certain about my skepticism. The fractions here in the details are debatable. Within the text of the paper, the author admits that the true ancestral populations are probably not represented by the model. But they are close. In most cases, the “Han” ancestry is probably indicative of the fact that the non-ANI component of South Asian ancestry is most closely related to the Onge, but is significantly different nonetheless.

The ratio of Iran_N and Steppe_EMBA is the key. Here is a selection from the paper:

Group Iran_N Steppe_EMBA Ratio
Jew_Cochin 0.53 0.23 2.27
Brahui 0.60 0.30 1.98
Kharia 0.13 0.07 1.97
Balochi 0.57 0.32 1.75
Mala 0.23 0.18 1.25
Vishwabrahmin 0.25 0.20 1.21
GujaratiD 0.29 0.28 1.04
Sindhi 0.38 0.38 1.00
Bengali 0.22 0.25 0.91
Pathan 0.36 0.45 0.81
Punjabi 0.24 0.33 0.72
GujaratiB 0.27 0.38 0.72
Lodhi 0.21 0.29 0.72
Burusho 0.27 0.43 0.64
GujaratiC 0.23 0.37 0.61
Kalash 0.29 0.50 0.58
GujaratiA 0.26 0.46 0.57
Brahmin_Tiwari 0.23 0.44 0.51

Any way you slice it, a group like the Tiwari Brahmins of Northern India have more Onge-like ancestry than most of the groups in Pakistan. But also observe that the ratio toward Steppe_EMBA is more skewed in them than among even Pathans or Kalash.  The Lodhi, a non-upper caste population from Uttar Pradesh in north-central South Asia are more skewed toward Steppe_EMBA than Pathans.

It is important for me to reiterate that the key is to focus on ratios and not exact percentages. Though the Steppe_EMBA fraction did strike me as high, glimmers of these sorts of results were evident in model-based clustering approaches as early as 2010. The population in the list above most skewed toward Iran_N are Cochin Jews. This group has known Middle Eastern ancestry. But next on the list are Brahui, a Dravidian speaking group in Pakistan. There is a north-south cline within Pakistan, with northern populations (Burusho) being skewed toward Steppe_EMBA and southern ones (Sindhi) being skewed toward Iran_N. Additionally, Iranian groups such as Pathans and Baloch likely have had some continuous gene flow with Middle Eastern groups, probably inflating their Iran_N.

Trends I see in the data:

  1. There is a north-south cline within Pakistan with Steppe_EMBA vs. Iran_N
  2. There is a north-south cline within South Asia with Steppe_EMBA vs. Iran_N
  3. There is caste stratification within regions between Steppe_EMBA vs. Iran_N
  4. Though not clear in this table, there are strong suggestions that Indo-European speaking groups tend to be enriched in Steppe_EMBA, all things equal (e.g., the Bengalis in the 1000 Genomes look a lot like the middle-caste Telugus in the 1000 Genomes when you remove the East Asian ancestry…except for a noticeable small fraction of a component which I think points to Indo-European ancestry)

What does this mean in terms of a model of the settlement of South Asian over the past 4,000 years? One conclusion I have come to is that Dravidian speaking groups are not the aboriginal peoples of the subcontinent. Rather, their settlement across much of South Asia is very recent. Almost as recent as Indo-Aryan habitation. In First Farmers the archaeologist Peter Bellwood proposed this model, whereby Indo-Aryans and Dravidians both expanded across South Asia concurrently. Though I think elements of Bellwood’s model that are incorrect, it’s far more correct in my opinion than I believed when I first encountered it.

Why do I believe this?

  1. The Neolithic begins in South India in 3000 BC.
  2. Sri Lanka is Indo-European speaking
  3. The Dravidian languages of South India don’t seem particularly diverged from each other
  4. There is ancestry/caste stratification in South India even excluding Brahmins (e.g., Reddys and Naidus in Andhra Pradesh look somewhat different from Dalits and tribals)
  5. Some scholars claim that there isn’t a Dravidian substrate in the Gangetic plain
  6. R1a1a-Z93, almost certainly associated with Indo-Aryans, is found in South Indian tribal populations
  7. Using LD-based methods researchers are rather sure that the last admixture events between ANI and ASI (“Ancestral South Indians”) populations occurred around ~4,000 years ago

Here is my revised model as succinctly as I can outline it. The northwest fringes of South Asia, today Pakistan, and later to be the home of the Indus Valley Civilization (IVC), was populated by a mix of indigenous populations, a form of ASI, when West Asian agriculturalists arrived ~9,000 years ago from what is today Iran. These were the Iran_N or “eastern farmer” groups. The West Asian agricultural toolkit was serviceable in northwestern South Asia for reasons of climate and ecology, but could not expand further east and south for thousands of years.

There is where the first admixture occurred that led to a population was mixed between ANI and ASI. These people lacked Steppe_EMBA. They were pre-Indo-European. They were almost certainly not all Dravidian speaking. The Burusho people of northern Pakistan, for example, speak a language isolate (in India proper you have Nihali and Kusunda)

By ~3000 BC this proto-South Asian (in a modern sense) population began to expand, while the IVC matured and waxed. Eventually, the IVC waned, fragmented, and disappeared.

Around ~2000 BC, or perhaps somewhat later, Indo-Aryans arrive in South Asia. The situation at this stage in not one of a primordial and static Dravidian India, on which Indo-Aryans place themselves on top. Rather, it’s a dynamic one as the collapse of the IVC has opened up a disordered power vacuum, and a reconfiguration of cultural and sociopolitical alliances.

In the paper above the author alludes to the pervasiveness of both Iran_N and Steppe_EMBA ancestry in South Asia, including in South India. “Indo-European” Y chromosomal lineages are also found among many South Indian groups, albeit at attenuated proportions region-wide. In Peter Turchin’s formulation, I believe that “Indo-Aryan” and “Dravidian” identities became meta-ethnic coalitions in the post-IVC world. Genetically the two groups are different, on average. But some Dravidian populations assimilated and integrated Indo-Aryan tribes and bands, while Indo-Aryans as newcomers assimilated many Dravidian populations.

The reason that the ratio of Iran_N to Steppe_EMBA does not decline monotonically as one goes from west to east along North Indian plain is that Indo-Aryans were not expanding into a Dravidian India.  Dravidian India was expanding only somewhat ahead of Indo-Aryan India, and in some places not all at all. In the northwest fringe of South Asia there had long been a settled population of peasants with West Asian ancestry with Iran_N affinities. In contrast to the east the landscape was populated by nomadic tribal populations with ASI affinities. North Indian Brahmins may have more Steppe_EMBA than some populations in Pakistan and more ASI because they descend from Indo-Aryan groups who absorbed indigenous ASI populations as they expanded across the landscape.

Dravidian groups as they expanded also assimilated indigenous populations. This explains some groups with very high fractions of ASI. Their ASI ancestry is a compound, of an old admixture in Northwest India, and also later assimilation in South India. The presence of R1a1a-Z93 in these populations reflects the integration of some originally Indo-Aryan groups into the expanding Dravidian wavefront.

Where does this leave us?

  1. The Indo-Aryan vs. Dravidian dichotomy is not one of newcomers vs. aboriginals. It is of two different sociocultural configurations which came into their current shape in the waning days of the IVC. That is, it is less than 4,000 years old
  2. The two populations were clearly interacting closely around the time of the collapse and disintegration of the IVC and post-IVC societies. There has been gene flow between the two
  3. ~4000 years ago ANI and ASI populations existed in their “pure” form, but that is because ASI aboriginals still existed to the south and east of the IVC, while Indo-Aryans were a new intrusive presence in the Indian subcontinent