The Pith: Afro-Indians are mostly African, with a substantial Indian minority ancestry. The latter is disproportionately female mediated. It also seems that that ancestry is more northwest Indian, and that natural selection has been operating upon them outside of the African environment.
Along the western coast of South Asia, from Makran in southwest Pakistan, down to the Konkan coast of southwest Iindia, there are isolated communities of Afro-Indians. They are called Siddis or Habshi. Their African origin is clear in their physical appearance, as well as aspects of their folk customs which tie them back to Sub-Saharan African. Nevertheless, they have assimilated to many Indian cultural traits. They generally speak the local language, and practice Islam, Hinduism, or Roman Catholic Christianity (in that order in proportion).
How and why did the Siddis arrive in India? The earliest date for their arrival almost certainly must be bounded by the period when Indo-Islamic polities rose to prominence in the early second millennium. The cosmopolitan melange of the armies of the Muslim warlords included diverse groups of Africans, some of whom took power, and established their own self-conscious Afro-Indian dynasties, set apart from the Turkish, Afghan, Persian, and Arab inflected statelets. Were these the sources of the modern Siddi communities? The oral history of the Siddi of the western coast of South Asia suggests not. In fact the geographical concentration of these Afro-Indian tribes along the Arabian sea fringe is indicative of different historical actors: the Portuguese. In much of Asia, out to China, the role of Africans was very different from that in the New World. They were objects purchased as for elite consumption, not production. They served at court, guarded the harem, etc. Lowland Asia had no need for imported labor, as there was human stock aplenty. Whereas in much of the New World black African slaves were critical cogs in the capitalist system of production, in Asia, as in the Arab world outside of a few areas such as southern Iraq, they were signals of luxurious consumption by the high and mighty (this was in vogue at European courts for a period as well).
Two new papers published yesterday in the American Journal of Human Genetics examine the genetics of the Siddi of India with an eye toward elucidating the details of their historical ethnogenesis. Though the papers overlap to a great extent, there are subtle differences which result in complementation. Shah et al. uses a far thicker set of markers, while Narang et al. look at many more populations, but due to removing SNPs which don’t span their populations the marker set is much thinner. Let’s review the papers in turn.
Two years ago Reconstructing Indian Genetic History reframed how we should view South Asian historical genomics. In short, Indians can be viewed as a hybrid between a West Eurasian group, “Ancestral North Indians” (ANI) and a very different group, “Ancestral South Indians” (ASI), which had distant connections to West and East Eurasians. At least to a first approximation. Last fall I posted on a new paper which surveyed the Austro-Asiatic speaking peoples of India, and concluded that they were exogenous to the subcontinent. This is an interesting point. Prehistoric treatments of South Asia often use linguistic terms to denote putative ancient populations. One model is that first it was the Munda, the most ancient Austro-Asiatics. Then the Dravidians. And finally the Indo-Aryans. These genetic data imply that the Munda arrived after the initial ANI-ASI synthesis. The Munda people of India can be thought of as ANI-ASI, with an overlay of East Eurasian ancestry.
Zack Ajmal’s K = 11 ADMIXTURE run has highlighted some further issues. He has a set of Austro-Asiatic samples, as well as a host of Indo-Aryan and Dravidian speaking populations. I now believe we can now further clarify and refine our model of the peopling of India. Here it is:
1) ASI, circa ~10,000 years BP
2) ANI enters the subcontinent from the northwest, synthesis with ASI
3) The ancestors of the Munda enter from the northeast, synthesis with ANI + ASI in their region
4) A subsequent group of West Eurasians, related to the ANI, so I will term them ANI2, enters from the northwest and overlays the ANI + ASI synthesis. In the northeast quadrant of the subcontinent this group marginalizes the Munda people, who are either assimilated or escape to more remote locations. I believe that ANI2 is likely the Indo-Europeans, but it may be Dravidians as well
5) A second group of Austro-Asiatic peoples enters from the northeast, and synthesizes with the AN2 + ANI + ASI. In some regions they are absorbed (Assam), but in other regions they are culturally dominant (Meghalaya)
Below are two plots which illustrate where I’m coming from. The “S Asian” component from K = 11 above seems to overlap, but is not identical to, ANI. The “Onge” component plays a similar role with ASI. The “SW Asian” and “European” elements are pretty straightforward. They’re very closely related to the “S Asian” one, but they do separate from it. Their relationship to distant non-Indian groups as well as a gradient toward the northwest suggests to me a more recent arrival of this element.
Zack Ajmal now has over 50 participants in the Harappa Ancestry Project. This does not include the Pakistani populations in the HGDP, the HapMap Gujaratis, the Indians from the SVGP. Nevertheless, all these samples still barely cover vast heart of South Asia, the Indo-Gangetic plain. Here is the provenance of the submitted samples Zack has so far:
Again, note the underrepresentation of two of India’s most populous states, Uttar Pradesh, ~200 million, and Bihar, ~100 million. Nevertheless, there are already some interesting yields from the project. Below I’ve reedited Zack’s static images (though go to his website for something more dynamic) with the labels of individuals. I’ve highlighted myself and my parents with the red pointers.
Zack has been posting his data sources, as well as how he filtered and formatted them, all this week. I assume that the first wave of results will be online soon. As of yesterday, this is what he had (I know he got some more today):
– Punjab 7
– Bengal 1
– Bihar 1
– Tamil 5
– Karnataka 1
– Anglo-Indian 1
– Roma 1
– Iran 3
Whole swaths of north-central India are missing. I am hopeful that more people will join in after the first wave of results are put out there. But, from what I have discussed with Zack it looks plausible that the very first wave will have a richer set of results because of the necessity of preliminary steps. So there’s some benefit in getting early. It’s really ridiculous to have literally 1 sample representing the 300 million people of Uttar Pradesh and Bihar. That’s 25% of South Asians represented by one person. I’ve gotten a commitment from one friend who was born U.P. to give his data up once it comes in, but there have to be others out there. (the Bengali N should go up to 2 when I swap my parents in for me)
The public data sources have Gujaratis, Tamils, Pakistanis (Punjabis, Pathans, Sindhis), and some South Indian groups (Tamil and Telugu). This leaves a blank spot on the North Indian plain.
Here’s the brief for the project again.
Quick review. In the 19th century once the idea that humans were derived from non-human ancestral species was injected into the bloodstream of the intellectual classes there was an immediate debate as to the location of the proto-human homeland; the Urheimat of us all. Charles Darwin favored Africa, but in many ways this ran against the cultural grain. The theory of evolution was birthed before the highest tide of the age of white supremacy and European hegemony, and Darwin’s model had to swim against the conviction that Africans were the most primitive of the colored races. After the waning of the ideological edifice of white supremacy, and the shock it received during and after World War II, the debates as to the origin of humanity still remained contentious and followed the same outlines (though without the charged normative inferences). But as the decades wore on many more researchers began to believe that Darwin was correct, and that the origin of humanity lay in the African continent. First, the deep origin of the human lineage in Africa was accepted, but eventually a more recent expansion out of Africa was argued for by one school. The turning point in these academic disputes was the popularization of the “mitochondrial Eve” theory of the 1980s.
What some paleontologists had long argued, that anatomically modern humans have their locus of origin in Africa, was supported now by research from genetics which indicated that Africans were the most basal clade of humans on a continental scale, so that non-Africans could be conceived of as a subset of Africans. From this originates the chestnut of wisdom that Africans have more genetic diversity than all other human populations combined. By the year 2000 one could say that the “Out of Africa” triumphalism had proceeded to the point where an almost exterminationist model had taken hold when it came to the relationships of anatomically modern H. sapiens, and other groups which had evolved outside of Africa over the past million or so years, such as the Neandertals.
But the theoretical dichotomies were too coarse and absolute as it turns out. A division between multiregionalist phyletic gradualism, where H. sapiens evolved out of its hominin ancestors concurrently on a world wide scale, and a model of rapid expansion of one tribe in Africa to replace all others in totality, may have been warranted in the age of classical genetics and a morphometric analysis, but now we can look at the raw genomic material in a more fine-grained fashion. In fact, we can now look at the genomic patterns of variation among extinct hominins! Though there have long been hints that the expansion-and-replacement paradigm was too extreme from the genetic and morphological data, with the publication last spring in Science of a paper which made the claim for admixture between Neandertals and non-Africans in the range of 1-4% in all non-African groups based on a comparison of Neandertal and modern human genetic variation, one can dismiss absolutist expansion-and-replacement as self-evidently true orthodoxy. But one orthodoxy has no given way to another, and the shock to the old models presented by the data has not resulted in the coalescence of new robust paradigms. We live in a time of scientific troubles, so to speak.
I have put up a few posts warning readers to be careful of confusing PCA plots with real genetic variation. PCA plots are just ways to capture variation in large data sets and extract out the independent dimensions. Its great at detecting population substructure because the largest components of variation often track between population differences, which consist of sets of correlated allele frequencies. Remeber that PCA plots usually are constructed from the two largest dimensions of variation, so they will be drawn from just these correlated allele frequency differences between populations which emerge from historical separation and evolutionary events. Observe that African Americans are distributed along an axis between Europeans and West Africans. Since we know that these are the two parental populations this makes total sense; the between population differences (e.g., SLC24A5 and Duffy) are the raw material from which independent dimensions can pop out. But on a finer scale one has to be cautious because the distribution of elements on the plot as a function of principal components is sensitive to the variation you input to generate the dimensions in the first place.
I can give you a concrete example: me. I showed you my 23andMe ancestry painting yesterday. I didn’t show you my position on the HGDP data set because I’ve shared genes with others and I don’t want to take the step of displaying other peoples’ genetic data, even if at a remove. But, I have reedited some “demo” screenshots and placed where I am on the plot to illustrate what I’m talking about above. The first shot is my position on the two-dimensional plot of first and second principal components of genetic variation from the HGDP data set.
Dienekes has a post up where he highlights the fact that the recent paper on South Asian metabolic diseases has a figure which elucidates population structure within the region. Accounting for structure is important for genome-wide associations since you might get a spurious correlations if trait value/disease frequency is simply tracking cryptic population variation. Dienekes says:
The existence of two clusters is kind of obvious, while their interpretation is not as dots of the same color appear in both clusters: a placement of these individuals in a global context might have been useful here. Things are clearer at the top cluster which shows a clear gradient anchored by Punjabi Sikh and Hindu Tamils on either end.
Also of interest is the group of isolated Muslim/Christian individuals on the left which deviate strongly from the mainstream; these probably represent exogenous elements that don’t resembe the bulk of the Indian population.
The second issue is easily addressed. The Christian outliers are both give English as their native language. That suggests to me that they’re Anglo-Indian, a community of mixed South Asian and European origin. South Asian Muslims are overwhelmingly of indigenous origin. But, a minority of the Muslim elite are West Asian, or have substantial West Asian ancestry, as is evident by the fact that they look white. Benazir Bhutto’s mother was of Kurdish and Persian ethnic background (her family was from Esfahan in Iran). I’ve reedited the religious & linguistic PC plots to fit onto the screen.