Razib Khan One-stop-shopping for all of my content

May 2, 2021

Men of the North

Filed under: Genetics,Historical Population Genetics — Razib Khan @ 11:43 pm
RegionI1I2*/I2aI2bR1aR1bGJ2J*/J1E1b1bTQN
Russia510.504661302.51.51.523
Lithuania66138500010.50.542
Latvia611401200.500.50.50.538
Estonia1530.53280102.53.50.534
Finland2800.553.50000.50061.5
Sweden371.53.51621.512.50302.57
Norway31.504.525.53210.501012.5
Denmark3425.515332.5302.5011

(Y chromosomal haplogroups)

A few weeks ago I saw the Y chromosomal haplogroup group distribution in Finland and Sweden. I’d know this disjunction for a while, but it really struck me. I got the numbers above from Eupedia, but you can find them elsewhere. Most of you probably know that Finland has a high fraction of N (they keep changing the nomenclature, so I’ll leave the number off). What’s curious to me is how low the fraction of N in the rest of Scandinavia is. Much of the N we see in Sweden may even be historical era migration of Finns into Sweden when the two nations were in political union (Finland was basically a Swedish colony).  Another notable fact is that N is very common among Baltic people, whether Finnic in the language (Estonian) or Indo-European (Latvian and Lithuanian).

Another strange thing is that while the Indo-European lineages of R1 are both at very low frequency in Finland, I1, which is common to the west in Scandinavia, is not. The latest ancient DNA makes it clear that Finnic languages seem to have arrived in the Baltic in the period between 1000 and 500 BC. Before then Corded Ware/Battle Axe people seem to have been dominant in the East Baltic. These people usually carried Y chromosome R1a.

The fact that N is so high in the Baltic nations shows that newcomers arrived, and in the northern region language shifted happened, but in the south, it did not. Meanwhile, further north in Finland almost all the R1a lineages disappeared. Not so with I1. There are all sorts of tortured explanations for this pattern, so I won’t offer one.

Genome-wide the Finns aren’t that different. The largest proportion of their ancestry is still Yamnaya/steppe:

I only post this to illustrate how strong “male-mediated” dynamics can be. The proportion of Siberian ancestry in Finns is rather low, but > 50% of their Y chromosomes are N. I think it is plausible that one of the reasons for the massive reduction in R1 in Finland might be due to climate change and massive population collapse among the Battle Axe people of southern Finland, and the later arrival of Siberians.

March 16, 2021

How the Irish became white

The Welsh writer and historian Norman Davies reflects in his magisterial book The Isles: A History that what sets the Irish apart from the British, and in particular the English, is that they retained their mythology. By this, Davies alludes to the fact that the Irish became Christian gradually and through a process of local adoption, rather than as an external shock, cajoled by foreign missionaries or coerced by their ambitious monarch. When St Patrick converted the Irish — an event celebrated today by millions around the world — it was a remarkably peaceful transition that left the country’s ancient culture alive.

Due to the gradual nature of transformation, the pagan lore of Ireland was recorded by monks who saw their work as a way to preserve and continue the legacy of their own native culture. J. R. R. Tolkien famously created the world of the Lord of the Rings so that the Anglo-Saxons could have their own mythos. The Irish have no need of such creative endeavours, since the tales of Cú Chulainn and Fionn mac Cumhaill have wrapped within them the pathos of Túrin Turambar and Achilles. The Lebor Gabála Érenn, “Book of Invasions,” more than matches The Silmarillion.

These are enduring myths that have been preserved in Irish culture — but what if I told you that the legend and myth are rooted in reality? That the Irish oral tales preserved by monks in the 8th century AD are echoes of events from thousands of years in the past? Far afield from folklore and oral history, the new science of ancient DNA is putting concrete flesh and bones upon the veiled prehistory in which Irish myth is rooted.

The story goes back to the last Ice Age when Ireland was mostly covered in ice and, like Britain, uninhabited. As the ice sheets retreated 10,000 years ago, the island began to be recolonised by Mesolithic hunter-gatherers arriving from the continent, venturing into an empty landscape. These people had migrated out of southern Europe, following the fauna and flora north as the continent warmed up, and were related to people then found in Britain, France and Spain.

They looked strikingly different from today’s Europeans. Genetic analysis indicates that their skin was dark, as was their hair — but their eyes were likely blue. The genetic difference between these people and later Europeans is similar to that between modern Europeans and Chinese. This world of hunter-gatherers persisted for thousands of years, as small bands clung to the western edge of Europe, practising a lifestyle with roots in the deepest Palaeolithic, eating a protein-rich diet of horses, bison, aurochs and red deer.

All this ended with the arrival of farming. A generation ago there were roiling debates among archaeologists as to whether agriculture came to northern Europe through cultural diffusion or migration, a debate similar to that which had raged about whether the Anglo-Saxons had conquered Britain in large numbers. Ancient DNA has now definitively answered this question, and we know that the first farmers in Europe descended from Anatolian migrants who swept in from the Near East. They spread across the continent rapidly around five millennia before Christ, mixing only minimally with the native hunter-gatherers, who were thin on the ground and could offer little resistance; although hunter-gatherer peoples had healthier diets and probably lived longer, farming societies could feed vastly more people and so overwhelmed their neighbours.

These newcomers arrived in Ireland around 4500 BC, many thousands of years after they had established themselves in southern Europe. They brought with them their grains, sheep and cattle, but many of the elements of Irish agricultural life were already in place and had been for over 1,500 years, conditioned by the local ecology and climatic regime. Though the new people mixed with the native hunter-gatherers, their own language and culture came to dominate, while ancient DNA indicates that the hunter-gatherers were very small in number. There were simply many more of the newcomers.

What few Mesolithic hunter-gatherers there were retreated and were absorbed in the advancing human wave of farmers. For many generations the two groups would probably have been easy to distinguish: the newcomers were light-skinned and dark-eyed, while the hunter-gatherers were dark-skinned and light-eyed. Ancient DNA suggests dark-skinned people persisted long after the arrival of farmers in pockets and corners of the island, in more remote and mountainous regions of the country. Who knows if some of the Irish legends of different people and races did not emerge from these contacts?

Curiously, the closest modern people genetically to the Neolithic farmers of Ireland are the Sardinians. But this is not surprising considering that most of the ancestry of the first Irish farmers seems to have derived from the Cardial Culture of Spain (itself the product of a rapid expansion of farmers from the eastern Mediterranean). While the Mesolithic hunter-gatherers were dark-skinned but light-eyed, the farmers likely resembled many modern southern Europeans, with dark hair and eyes, and lighter skin. The contact between two such distinct groups, almost certainly speaking unintelligible languages, physically so different and practising contrasting lifestyles, almost certainly fed into later legends and myths.

The arrival of farmers in Ireland led to a transformation of the material culture. These Neolithic people were responsible for the construction of great stone megaliths, a practice that arose and spread along Europe’s Atlantic shores, seen with Stonehenge in Britain and the Carnac Stones of Brittany. They also built monuments such as the Newgrange passage tomb, one of the most ornate and beautiful pieces of prehistoric stonework in the world. These various stoneworks and tombs that litter the landscape of Ireland have shaped the Irish sense of their past, and loom large in legends of faeries and ancient peoples long lost and faded into the mists of time.

But the time of the Stone Age farmers ended in due course, and in 2500 BC the first of many waves of migrants swept west onto the island. Called the “Beaker Culture“, they were descendants of people who had left modern Germany, and earlier still had their origins on the Eurasian steppe. Genetically, the Beaker People resemble the modern-day inhabitants of the island and physically, too, these newcomers appeared rather like the fair-skinned and often fair-haired modern Irish. Though many of the traits now associated with the Irish, such as lactose tolerance, were not present in full, the roots of the genetic character of the modern people date to this period.

One hypothesis holds that it was the Beaker People who brought the Celtic languages, establishing the cultural patterns that persisted down to the Iron Age and contact with the Romans. If this is true, then the myths that were recorded by the Irish monks in the 7th and 8th centuries AD may date to the period of contact and conflict between the Neolithic tribes and the Beaker Culture. Could it be that the Tuatha De Danann, a mythologic race of supernatural beings mentioned in Irish legend, and vanquished by the ancestors of the Gaels, were actually the builders of the megaliths?

Recently, a group of Irish scientists has cataloged genetic remains from a host of locations across Neolithic Ireland. They discovered that a high-status man buried at Newgrange 5,000 years ago appeared to have been part of an important prehistoric dynasty, and was also the product of incest. The researchers point out that it is often a feature of highly-stratified Neolithic societies that royal families will practise incest, yet it is intriguing that the early historical Irish had legends of incest at the site of Newgrange, a story that must have survived thousands of years until Christianisation and literacy.

The local people in fact called the mound the “Hill of Sin”, and there were legends of incestuous activities between a ruler who built monuments and his sister in order to maintain the cycle of the sun. And so the existence of an individual who is quite inbred, and clearly of elite status, buried 3,000 years before these stories were written down is quite extraordinary.

Is it possible that Irish folklore was able to record an essentially truthful story over so many generations? The mythic legends of Ireland date to the time of Christ, over 2,000 years after the arrival of the Beaker Culture, and their likely conflict with the last of the Stone Age farmers who built the monuments they inherited. Preservation of such ancient motifs seems implausible. But note that Indian mythology preserves elements that are resonant with Greek myth, such as the divine twins — this, despite the two cultures being separated by thousands of years since their last common ancestor, suggesting a story that had been maintained for generation upon generation.

Davies, among other historians, highlights the Irish preservation of their mythic cycle as a matter of curiosity, and perhaps an edifying fact. But the detail and fidelity of the Irish in setting down their oral history may actually benefit future generations, as ancient DNA is telling us in exquisite detail the demographic processes by which the Irish people came to be the way they are. The true book of invasions is being written by unraveling DNA, and the existence of an oral record of the deep past may be essential for future scholars in understanding life in the islands before the Romans brought it into the light of history.

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February 26, 2021

Steppe lineages in northern Pakisan

Filed under: Genetics — Razib Khan @ 2:38 pm

This is not the most important paper, but it is a contribution: Complete mitogenomes document substantial genetic contribution from the Eurasian Steppe into northern Pakistani Indo-Iranian speakers. Abstract:

To elucidate whether Bronze Age population dispersals from the Eurasian Steppe to South Asia contributed to the gene pool of Indo-Iranian-speaking groups, we analyzed 19,568 mitochondrial DNA (mtDNA) sequences from northern Pakistani and surrounding populations, including 213 newly generated mitochondrial genomes (mitogenomes) from Iranian and Dardic groups, both speakers from the ancient Indo-Iranian branch in northern Pakistan. Our results showed that 23% of mtDNA lineages with west Eurasian origin arose in situ in northern Pakistan since ~5000 years ago (kya), a time depth very close to the documented Indo-European dispersals into South Asia during the Bronze Age. Together with ancient mitogenomes from western Eurasia since the Neolithic, we identified five haplogroups (~8.4% of maternal gene pool) with roots in the Steppe region and subbranches arising (age ~5–2 kya old) in northern Pakistan as genetic legacies of Indo-Iranian speakers. Some of these haplogroups, such as W3a1b that have been found in the ancient samples from the late Bronze Age to the Iron Age period individuals of Swat Valley northern Pakistan, even have sub-lineages (age ~4 kya old) in the southern subcontinent, consistent with the southward spread of Indo-Iranian languages. By showing that substantial genetic components of Indo-Iranian speakers in northern Pakistan can be traced to Bronze Age in the Steppe region, our study suggests a demographic link with the spread of Indo-Iranian languages, and further highlights the corridor role of northern Pakistan in the southward dispersal of Indo-Iranian-speaking groups.

Don’t focus on the percentages too much. Rather, focus on the coalescence estimate. Basically, that indicates diversification and demographic expansion. The presence in the southern subcontinent is indicative of the fact that “steppe” ancestry and cultural influence extends far beyond the distribution of modern Indo-Aryan languages. R1a we know, as it is found in adivasis. And low fractions of steppe are found in most South Indian groups (but not all).

February 9, 2021

The Genetics of India Cloubhouse Event – Friday 9 PM CDT

Filed under: Genetics — Razib Khan @ 11:55 pm

I am hosting a Clubhouse room this Friday, 9 PM CDT (8:30 AM in India on Saturday). The topic will be the genetics of India, and I’ll be talking about my two posts on Substack:

The Stark Truth About Aryans

The Stark Truth About Humans

It’s basically going to be an interactive discussion. My friend David Mittelman will help me moderate (probably others too).

You have to have a Clubhouse account (iPhone only). If you want to follow me on Clubhouse, I’m @razibkhan just like on Twitter.

January 24, 2021

Why Scythians, Sakas, and Kushanas, are NOT the source of “steppe” ancestry

Filed under: Genetics — Razib Khan @ 7:06 pm

This is a common question/assertion in the comments pretty much every other week: why couldn’t the documented incursions of nomadic people in the first millennium A.D. be responsible for the steppe ancestry? There is actually a good explanation in The formation of human populations in South and Central Asia, so I’ll quote it:

By the Late Bronze Age, ESHG-related admixture became ubiquitous, as documented by our time transect from Kazakhstan
and ancient DNA data from the Iron Age and from later periods in Turan and the Central Steppe, including Scythians, Sarmatians, Kushans, and Huns (29, 52). Thus, these first millennium BCE to first millennium CE archaeological cultures with documented cultural and political impacts on South Asia cannot be important sources for the Steppe pastoralist–related ancestry widespread in South Asia today (because present-day South Asians have too little East Asian–related ancestry to be consistent with deriving from these groups), providing an example of how genetic data can rule out scenarios that are plausible on the basis of the archaeological and historical evidence alone (13) (fig. S52). Instead, our analysis shows that the only plausible source for the Steppe ancestry is Steppe Middle to Late Bronze Age groups, who not only fit as a source for South Asia but who we also document as having spread into Turan and mixed with BMAC-related individuals at sites in Kazakhstan in this period. Taken together, these results identify a narrow time window (first half of the second millennium BCE) when the Steppe ancestry that is widespread today in South Asia must have arrived.

There is now a large database of Scythian, etc., ancient DNA, thanks to the preservation conditions on the Eurasian steppe. Most of their ancestry derives from the same broad group as the Andronovo horizon of which the Sintashta were part. But, unlike the earlier steppe populations, these groups are highly variable in ancestry, as well as usually having substantial minority East Asian components. The Indian groups with a lot of steppes, such as Jatts and Northern Brahmins, lack this.

There are two objections. The weaker one is that they didn’t have statistical power to detect the admixture. I haven’t run simulations, but I’m sure they have. If you have Jatts who are perhaps more than 30% steppe they would have detected trace East Asian (as you can find in many Muslim individuals from Pakistan).

The stronger objection is that there is unsampled structure on the steppe, and groups without East Asian admixture that are direct descendants of the Sintashta without dilution. This is not entirely unreasonable or implausible, though at this point I’d say this is unlikely for two reasons:

  1. Central Eurasia is pretty well sampled due to interest and conditions
  2. The steppe ancestry in South Asia is pretty widespread. Hard to imagine it percolating so far in 1,500 years

Also, the statistical tests I’ve done show Bengalis got East Asian admixture 1,500 years ago. 10-20% of the ancestry. The steppe percentage in Bengalis is 10-15%. But I never get any hits using older less sensitive methods of admixture. That means that it has to be way older a mix than 500 AD.

January 19, 2021

Stark Truth About Aryans: a story of India (part 1)

Filed under: Genetics — Razib Khan @ 1:38 pm

My Substack piece is up, Stark Truth About Aryans: a story of India. I’m pretty proud of this, as it wasn’t a single-sitting blog post, but something I worked over several times. Since it’s for paid subscribers I’ll post the first few paragraphs below, with an infographic that I think illustrates a lot of what’s going on.

Nearly one in four human beings lives in the Indian subcontinent. The region’s genetic and cultural diversity are unparalleled, spanning a vast spectrum from blue-eyed Muslim Kashmiris in the north who speak a tongue distantly related to English, to dark-skinned animistic Tamils in the far south whose language has no known relatives outside South Asia. Though Indian genetic research goes back decades, a legacy of P. C. Mahalanobis, the “father of Indian statistics,” only over the past twenty years has our understanding of the present genetic and physical variation and its roots expanded to a point where we can confidently trace the origins of South Asia’s bewildering diversity.

But science doesn’t exist in a vacuum. What the latest findings mean to Indians is still an issue subject to public debate, long after the data has been collected, reported, and analyzed. What does it mean to say that half the ancestors of modern Indians were related to the people of the Andaman Islands, and the other half to Europeans? Or that this synthesis of lineages occurred only within the last 10,000 years? Instead of being ancient and primal, a child of the  Pleistocene tens of thousands of years in the past, to be genetically Indian is to be younger than the pyramids of ancient Egypt. Indian diversity is the result of a massive mixing between various streams of humanity that occurred thousands of years ago to produce a mosaic with startlingly different shades and features. European pastoralists, farmers from the subcontinent’s northwest reaches, and hunter-gatherers who clung to the Indian-Ocean shores on the southern fringe of India, came together to stitch a complex tapestry.

Though the Indian press and public take a keen interest in how their genetic origins map onto the history of their storied civilization, interpretation of any empirical results is inevitably politically fraught. Geneticist David Reich recounts in his book Who We Are and How We Got Herealmost losing a 2009 paper right before publication due to its potential social and political ramifications. As Reconstructing Indian Population History was coming out, he had to navigate treacherous socio-political shoals. Indian collaborators made it clear that there were limits to what he could explicitly conclude because of cultural red lines. If Reich’s lab insisted on concluding that there was a massive migration of people into the subcontinent from the northwest, the Indian researchers would have to withdraw their cooperation, and therefore their essential data.

Reich’s work showed that all the nearly two billion people of Indian subcontinental origin emerged out of a fusion of two very distinct ancestral populations, which he termed “Ancestral North Indian” (ANI) and “Ancestral South Indian” (ASI). This amalgamation was then thought to have occurred only about 4,000 years ago. The closest contemporary relatives of the ASI are the indigenous people of the Andaman Islands, while the ANI are genetically similar to today’s Europeans and Middle Easterners.

Genetic

December 29, 2020

The massive Indian migration to Southeast Asian

Filed under: Genetics,Southeast Asia — Razib Khan @ 2:38 pm


Over at my other weblog I put up a post, Indian Ancestry In Southeast Asia Is Older Than Statistical Genetic Tests Suggest. If you look at two populations in Southeast Asia and find one has Indian ancestry you often can’t find the admixture older than 1000 A.D. (in peninsular Malaysia there is more recent intermarriage between Muslim Indians and Malays too). This seems far too recent. My explanation is simple: these dates reflect the assimilation of a hybrid Indian-Southeast Asian population across much of Southeast Asia. I have done the analyses myself, and in Cambodia, I get dates around 1000 A.D. Cambodia is not close to India and there isn’t evidence of a large Diaspora in recorded history. But, we know that Hinduism was a major influence in the region, and the Vietnamese Cham are still predominantly Hindu.

The kingdom of Funan, known mostly from Chinese accounts, flourished in Cambodia for the first five centuries of the common era or so. There is an inscription in Sanskrit from the region dated to the 5th century A.D. that refers to the moon of the Kauṇḍinya line (… kauṇḍi[n]ya[vaṅ]śaśaśinā …) and chief “of a realm wrested from the mud”. The text is in the Grantha script.

Further west, Dvaravati also had a strong Indic influence, no later than the 5th century A.D.

The genetic results indicate on the order of 10-20% of the ancestry of people in central Thailand is broadly Indian. This is not a trivial fraction. Who were these people? How early did they come?

On a minor editorial note, I’ll observe there is lots of discussion about possible Indian gene flow to the north and west (into Iran and Turan), but the data on Southeast Asia is clear and of greater magnitude. But there is far less discussion and exploration of this.

November 21, 2020

How much “steppe” ancestry is there in South Asia? (Indian subcontinent)

Filed under: Genetics,Sintashta,steppe — Razib Khan @ 5:46 pm

Since this question always comes up at some point, I decided to do a rough back-of-the-envelope calculation of the % steppe across the Indian subcontinent. The way I did it was by taking Pakistan, Bangladesh, and India, and estimating the average percentage from the caste breakdowns (e.g., UP is 20% “upper caste” and 20% “Dalit” and 60% neither, with fractions of steppe/Sintashta about 30%, 10%, and 15%, respectively).

So the final number I came back is that 14% of the ancestry in modern-day South Asia is from the steppe in the form of people descended from Sintashta pastoralists. That is about 220 million human beings worth. You can judge whether that’s significant or not. Additionally, it looks like closer to 20-25% of the Y chromosomes are derived from these people.

I’m not “showing my work” because I think no matter how you estimate it, you’ll get a number in this range. Perhaps 12%. Perhaps 16%. But what difference does that make?

October 28, 2020

The Evolutionary History of Man’s Best Friend Revealed

Filed under: Biology,Genetics,science,Science & Tech,Spotlight — Razib Khan @ 10:28 pm

Man and dog share a long history. In much of the world, a history as old as humanity. The latest genetic evidence now tells us that the emergence of the domestic dog lineage occurred soon after the human expansion out of Africa 50,000 years ago, in the depths of the last Ice Age. We came. We saw. And we befriended. This we knew, but now we can closely examine how. A paper out today in Science uses 27 ancient dog genomes from the past 11,000 years to construct an evolutionary history nearly as rich as that produced by human population geneticists over the last decade.

The authors found five lineages of ancient dogs that were present at the end of the last Ice Age. These were the dogs that interacted with human migrations during the rise of agriculture and the fall of civilizations to produce the riotous dog diversity that we know today. Familiar breeds like the Pekingese and the St. Bernard, as well as stray Asian village mutts, they’re all the products of a deep shared history, which has left its imprint on the genetic variation of modern dogs.

The dog-human relationship has always been special in several ways. Our canine companions are well adapted to us, in that they can engage with humans socially in a manner no other animal seems equipped to, not even our closest relatives, the chimpanzees. This is thanks to tens of thousands of years of co-evolution. In a very real sense the evolutionary niche of the domestic dog is the human mind. Wolves may have higher IQs, and yet there are 1,500–3,000 times more dogs alive today. Perhaps it was smarter to be friendly than just smart.

Dogs are the only domestic species who have been with us since the Pleistocene, which ended 11,500 years ago. Crops, cattle, and cats all came later, during the Holocene, after the Ice Age. When Siberians crossed over to the New World 15,000 years ago, they also brought their dogs. These dogs served many roles. Friend, beast of burden, and yes, even food.

The New World dogs, who were present when the Spaniards landed in 1492, are one of the five great Pleistocene lineages from which all modern dogs descend. The authors of the Science paper call them the Neolithic Levant, Mesolithic Karelia, Mesolithic Baikal, ancient America, and the New Guinea singing dog. The last group includes Australian dingos and represents the ancient expansion of our species and its companions into Southeast Asia. Karelia is a region bordering Finland and Russia, while the Baikal dogs come from the lake of that name in east-central Siberia. These samples are termed Mesolithic because they are more recent than the Ice Age (7–11,000 years ago), but the peoples in Karelia and Baikal were still hunter-gatherers at that time. In contrast, the Neolithic Levant dogs date to 7,000 years ago, and were companions to agricultural populations.

All modern dogs can be thought of as a mixture between these five early Holocene lineages. Despite their highly specific names, the Karelian and Baikal lineages represent huge geographic distributions, not just the sample sites. In this regard, they are analogous to modern humans. As David Reich has shown in his magisterial work, Who We Are and How We Got Here, the past 10,000 years have witnessed the merging of disparate hunter-gatherer tribes to form the peoples we see today, threaded together from distinct strands. This naturally leads us to the question: did human populations bring their particular dogs everywhere they went?

Comparing the relatedness of humans and relatedness of dogs, the authors of the Science paper found that there was a definite correlation. Phylogenetic trees of both dogs and humans exhibited strikingly similar shapes, indicating common underlying histories of separation and diversification. Analyses comparing the genetic data of dogs and humans revealed that they were tightly correlated. In fact, the relationship between the evolutionary trees for dogs and for humans is close enough that the tree for humans explains much of the pattern we see for dogs. There is an underlying structure which spans the two species, but also novel and curious deviations that alert us to particular stories.

Some of the stories are straightforward and to be expected. The dogs of ancient Europe are a mix of the Karelian and Neolithic Levant lineages. The latter clearly tagged along with the early farmers expanding out of the Near East. Further south in Europe, the dogs were more like those of the Neolithic Levant, while further north they were more like the Mesolithic Karelian dogs. American dogs were genetically closest to the Siberian dogs from Lake Baikal. Additionally, both these groups were closer to the dogs of Southeast Asia than they were to those of the Levant. This recapitulates the pattern for Eurasian peoples; East Asians are more closely related to Europeans than they are to Middle Eastern people.

But a few surprising findings confound our expectations. Four thousand years ago on the Volga steppe of Russia, on the cusp between Europe and Asia, the Srubna people flourished as part of a broad continuum of Indo-European peoples that stretched far to the west. While the Indo-European relatives of the Srubna came to dominate much of Europe, their dogs did not. The burials of early Indo-Europeans in Europe yield dogs whose heritage connected them to the Eurasian steppe and the Srubna, but this was a fleeting affair. The dogs of Northern Europe were not replaced by the dogs of the steppe, though the people largely were.

Instead, the heritage of the Srubna dog persists far to the east. Chinese dogs are mostly a mixture of ancient Southeast Asian dogs, related to the New Guinea singing dog and the dingo, plus the Srubna steppe dogs. While half the ancestry of modern Europeans comes from people related to the Srubna, very little of that ancestry is to be found in modern Chinese. And yet dogs are not the only evidence of the influence of steppe people in ancient China. In 1200 BC, chariots became prominent during the Bronze Age Shang dynasty, chariots whose design indicates borrowing from the Iranian peoples of Central Asia, direct descendants of the Srubna. It seems entirely likely that if chariots were brought to China during the Bronze Age, then so were dogs. While the dogs of the steppe found no welcome in Europe, the Chinese adopted them as their own.

It also sometimes happens that the migration of dogs runs counter to the migration of peoples. While humanity evolved in Africa and spread to the rest of the world through the Middle East, the origin of African domestic dogs reflects the reverse. Indigenous African breeds such as the Basenji descend from dogs of the Neolithic Levant. These dogs were likely brought south and west by the Near Eastern pastoralists who also brought cattle to the continent. Meanwhile, the ancient canine populations of the prehistoric Levant have been totally replaced by groups from Iran and Europe, so the primary legacy of the dogs that were associated with the first human cities is to be found today in Africa.

But perhaps the biggest twist of the great canine diasporas is that their recent history is obscured and dominated by the phenomenon which has overshadowed our own species’ history: the rise, expansion, and domination of the world by Europe.

Modern European dogs do not descend from the full diversity of the continent. Rather, almost all European breeds today derive from a population best represented by a 5,000-year-old sample discovered in southern Sweden, an individual who was equally descended from Karelian and Levantine populations. The authors could not pinpoint the time or place that this replacement of the ancient lineages occurred, but by the time of the rise of European colonialism this single population had swept aside all the other ancient populations of the continent.

Today the majority of the ancestry of dogs across the world is from this single European population that flourished 5,000 years ago. In the New World almost all the ancestry is now European, with the exception of the Arctic dogs. The venerable chihuahua has less than five percent of its ancestry from the dogs of the Aztecs. In Africa and the Middle East, most canine ancestry is now European, while European ancestry is often the majority in many Asian dog populations, and is substantial even in the dingo. Decolonization may have occurred politically, but the evolutionary history and genetic variation of dogs illustrate how powerful the hand of history can be and how long its shadow persists.

Ancient DNA is like a genetic time capsule. We’ve spent these first decades honing our extraordinary new tools as we examine our own ancient human ancestry. But ours is hardly the only species with a history worth unpacking in such fascinatingly granular detail. Rather, we can look forward to the recent history of life on Earth being thoroughly illuminated via the evidence left behind by hardy fragments of DNA from creatures long gone.

The past 40,000 years has seen the transformation of our planet’s ecosystems due to humans. We live in the “Anthropocene” because our species has reshaped the planet to our ends. The evolutionary history of the domestic dog reflects humanity’s first, and longest, bioengineering experiment. We took a pack of wolves and transmuted it into a new species to walk alongside us. We bred dogs to be useful tools, but eventually they became our most loyal allies. We owe it to them to understand how they came to be as they are. And in the process we’ll inevitably continue to learn about ourselves.

 

Razib Khan is a geneticist and director of science at the Insitome Institute. He has written for the New York Times, India Today, National Review Online, and runs two weblogs, Gene Expression and Brown Pundits. You can follow him on Twitter @razibkhan or visit his website here.

 

October 21, 2020

Tibeto-Burmans, Munda, and Bengalis

Filed under: Genetics — Razib Khan @ 10:58 am


I’m pretty sure I posted this Chaubey lab work as a preprint, but it’s now a published paper. For those who can’t understand the table, it illustrates a big difference between Tibeto-Burmans and Munda. The samples from Bangladesh look to be generic Bangladeshis, the 10% frequency for O2a seems to match the other data I’ve seen for East Bengalis.

This confirms that the East Asian admixture into Bengalis was not Munda. And, the Tibeto-Burmans of the nTibeortheast have no assimilated Munda ancestry. I think it does lend more credence to the idea that the Munda arrived in the Indian subcontinent across the Bay of Bengal, landing in Odisha, rather than from the northeast.

October 16, 2020

The rise of Indicus!

Filed under: Genetics — Razib Khan @ 12:00 am

A few years ago an ancient DNA paper on cattle was published, Ancient cattle genomics, origins, and rapid turnover in the Fertile Crescent. It’s a pretty good paper with interesting results. The paper confirmed pretty strikingly that there was a punctuated and massive expansion of indicus ancestry across the Near East between 3,500 and 4,000 years ago.

The interesting aspect of cattle is that there are really two species that intermix. Using mtDNA researchers estimate indicus and taurus diverged 300,000 to 2,000,000 years ago. But the main thing you have to remember is cattle generations are about 20% as long as human generations. So 300,000 cattle years are equivalent to 1.5 million human years. And, for technical reasons (smaller effective population size) one should probably assume mtDNA underestimates the divergence.

Ancient cattle from the Near East are all taurus. The PCA plot shows that most of the variance is on PC 1 which separates indicus and taurus (a secondary dimension is PC 2, between African and Near Eastern/European lineages). The figure at the top of this post shows that there is a massive jump in genome-wide indicus ancestry across the Near East between 2000 and 1500 BC. As the authors note this can’t be diffusion; the jump is too sudden and sweeping.

So what happened during this period? As noted in the paper: Bronze Age civilization almost collapsed around ~2000 BC. More concretely, after 2000 BC is when we see evidence of Indo-Europeans in the Near East. The Indo-Aryan Mittani show up in Mesopotamia in ~1600 BC. The Indo-European Hittites, the Nesa, are known from Anatolia a bit earlier.

This is also the period that small, but detectable, levels of “steppe” ancestry show up in some ancient samples.

Before this paper, I would have leaned to the position that the Mittani Indo-Aryans migrated directly from the Sintashta homeland without much contact with Indian Indo-Aryans. These data are too suggestive of a widespread zone of expanding agro-pastoralists that existed between western South Asia and the Near East between 2000 BC and 1500 BC.

One of the things we know from the barbarian period during the Fall of Rome is that barbarian groups had strong channels of information flow. For example, a group of Saxons arrived with the Lombards in Italy in the second half of the 6th century. But, through various channels, these Saxon warriors learned that their co-ethnics had established dominance in what was to become England, and there are texts which allude to the reality that they decamped and crossed the Alps, presumably on the way to what was going to be England. The point here is that there was a “Saxon international.”

Aside from the Mittani the evidence of Indo-Aryans in the Near East is tenuous, though some of the Kassites of Babylonia may have had Indo-European affinities. There is not nearly as strong a genetic imprint of steppe in the Fertile Crescent as in Northwest India. The Hittites were very different from Indo-Aryans, who seem to have the closest relationship to the Slavic language family.

The indicus breed is adapted to tropical dry climates. It seems plausible that the Indo-Aryan international facilitated the spread of this breed in the centuries before 1500 BC.

October 7, 2020

Armenia, Azerbaijan, Turkey, and genetics

Filed under: Armenia,Genetics,Turkey — Razib Khan @ 12:18 am

Recently a few people have been asking me about Armenian, Turks, and genetics. Mostly because I’ve written about this topic before. Unless you’ve been asleep you know that there is a war going on in the Caucasus. Armenia and Azeribaijan are renewing their decades-long conflict, and bringing other nations into as the local powers choose sides. Unpleasant all around.

Who are the Armenians? The Turks? Azeris?

Azeris in particular are not well known in the West, but they’re kind of a big deal. The Iranian province of Azerbaijan has nearly as many people as the Republic of Azerbaijan, and there are more Azeris in Iran than in independent Azerbaijan itself. The leader of Iran has an ethnic Azeri father. Azeris are Turkic and have traditionally been dominant in Iran’s military. Before the Turkification of the region over the last 500 years, Azerbaijan was called Albania. The native language was Iranian and related to Persian.

Living in close proximity to each other, it is no surprise that the peoples of the region are genetically rather similar. That being said, there are notable differences.

While a few Armenians in my datasets have Russian admixture (they are likely F1 individuals who identify as Armenian), what is notable about Azeris is like Turks they exhibit a small but notable shift toward East Asians. This is almost certainly the consequence of Turkic ancestry. Though most of the ancestry of Azeri is pre-Turkic, Turkification occurred through assimilation of nomads with some East Asian ancestry.

The sample applies to Turks to the west of Armenia. In previous posts, I’ve had discussions about the nature of Turkish ancestry, but in general, I am convinced by those who argue that the non-Turkic component (which would include East Asian and Turanian) reflects the earlier pattern of variation; Greek in the west, Armenian and Kurdish in the east.

Conflicts, like we are seeing today, illustrates the power of ideas over relatedness. Armenians are Christians, albeit peculiar Oriental Orthodox Christians. Additionally, they continue to speak an ancient Indo-European language. This sets them against Turkic speaking Muslims to the west, and Turkic speaking Muslims to the east, though all the groups share deep common ancestry.

October 3, 2020

Population structure in West Bengal and Bangladesh

Filed under: Genetics — Razib Khan @ 4:19 pm

The Genomes Asia 100K has put their Indian paper out. It’s OK, and mostly focuses on the fact that Indians are enriched for inbreeding vis-a-vis other world populations. There are several layers to this. In some cases, as among South Indian Hindus and Muslims, there is cousin-marriage. But, in other cases, for example, Scheduled Castes and Scheduled Tribes, there seem to be extreme bottleneck effects due to delimited marriage networks. Finally, even among large population groups, such as Iyers, there seems to be some elevation of runs of homozygosity due to endogamy.

But that’s really not what I’m interested in. This preprint has a lot of Bengalis from Birbhum district in West Bengal of various castes. The UMAP (an advance over PCA in some ways) figures aren’t super informative, but you can see that their pooled sample recapitulates the Indian subcontinent. In fact, West Bengals on the whole are to the “west” of Bangladesh samples. Totally unsurprisingly.

The main reason I’m putting this post up is the UMAP plot below. It’s hard to read (they will clean it up for final publication), and I don’t know all the castes (I’m assuming “Nabasudra” is a typo). But some things that jump out

1) Bengali Brahmins are distinct.

2) Kayastha are generic West Bengalis.

3) Some of the West Bengal samples are in the Bangladesh (collected from Dhaka) distribution. These are probably descendants of Bangal migrants from the east.

4) Some groups are very distinct. That’s partly due to strong endogamy, and in the case of Santhals high East Asian ancestry (they’re Munda). Other groups are less distinct. The “Namasudra” seem to be two groups. One overlaps with the main Bengali cluster (slight bias toward Bangladeshis), while a second group is shifted toward Scheduled Castes.

I assume readers can make more heads or tails of this, as I don’t know much about caste in West Bengal (and yes, the figure is very badly labeled/colored; this is a preprint)

Addendum: Not comments about Jatts please. I will delete them.

September 3, 2020

Ancient Pakistanis were Hindu

Filed under: Genetics — Razib Khan @ 2:11 pm

Over at my other blog, Pakistani British Are Very Much Like Indians Genetically. The title doesn’t refer to genome-wide worldwide affinities. Rather, the preprint looks at British Pakistanis, and finds a pattern that is not going to surprise Indians: endogamy seems to have kicked in for these groups starting 1,500 to 2,000 years ago. This is exactly what you see in the Indian jati data. The similarity is pretty incredible, and to me is a strong rejection of the model that these groups were strongly anti-caste so on the margins of Indic civilization.

There is a second wave of endogamy though, dated from 150-500 years ago, roughly. I think this is likely Islamicization and adherence to cousin-marriage. These Pakistani groups seem to show the tendency of jati endogamy common among Hindus, and, cousin-marriage patterns of the Islamic world.

Finally, the reason I posted over on the other blog is that I think this might speak to the long-term trajectories of Bangladesh and Pakistan: Bangladesh is not in the same mold as Indo-Pak societies. The 1000 Genomes data indicate few runs of homozygosity and not much internal structure. That is, no jati endogamy, and, low levels of cousin-marriage.

If you believe Joe Henrich, this means good things for Bangladesh in the future… (vs. Pakistan)

(the Henrich podcast is already available for Patrons)

Pakistani British are very much like Indians genetically

Filed under: Biraderi,Genetics,Pakistan — Razib Khan @ 1:51 pm

I talked to Joe Henrich this week for The Insight about his book, The WEIRDest People in the World: How the West Became Psychologically Peculiar and Particularly Prosperous (the episode is going life next week). Obviously much of the discussion hinged around relatedness, kinship, and how that impacted the arc of history (we also talked about other issues, such as the status of the “big gods” debate, so most definitely tune in!).

So I was very curious when I saw a new preprint, Fine-scale population structure and demographic history of British Pakistanis:

Previous genetic and public health research in the Pakistani population has focused on the role of consanguinity in increasing recessive disease risk, but little is known about its recent population history or the effects of endogamy. Here, we investigate fine-scale population structure, history and consanguinity patterns using genetic and questionnaire data from >4,000 British Pakistani individuals, mostly with roots in Azad Kashmir and Punjab. We reveal strong recent population structure driven by the biraderi social stratification system. We find that all subgroups have had low effective population sizes (Ne) over the last 50 generations, with some showing a decrease in Ne 15-20 generations ago that has resulted in extensive identity-by-descent sharing and increased homozygosity. Using new theory, we show that the footprint of regions of homozygosity in the two largest subgroups is about twice that expected naively based on the self-reported consanguinity rates and the inferred historical Ne trajectory. These results demonstrate the impact of the cultural practices of endogamy and consanguinity on population structure and genomic diversity in British Pakistanis, and have important implications for medical genetic studies.

None of this is entirely surprising. The media in the UK has written about recessive disease load because of cousin-marriage amongst Pakistani Britons. But there are also things in the preprint that need to be made explicit. The “biraderi” social system is apparently a paternal lineage system in the northwest of the Indian subcontinent which transcends religion (i.e., it is present across the border in Indian Punjab). These are “tribal” or “clan” societies in a way that is not present across much of the Indian subcontinent. For example, my family is from eastern Bengal. Before the partition between India and Pakistan, the far northwest and northeast of the subcontinent had the highest proportions of Muslims. But that did not mean that the two regions were culturally very similar, explaining in part the war in 1971 that resulted in Bangladesh. In Bangladesh, biraderi is not known, and the rates of cousin-marriage are much lower than in Pakistan.

One of the things I immediately noticed in the 1000 Genomes data is that Bangladeshis exhibit a lot less structure and stratification than Indians and the samples from Pakistani Punjab. In many ways, the patterns in the Bangladeshi genomes resemble the type of patterns in non-South Asian genomes: an outbreeding population without much internal structure.

This is not typical in South Asia. Rather, Indian populations tend to have lots of differences between jati/caste groups due to endogamy. To my surprise, Pakistani samples from Lahore were similar, though I attributed some of that to the migration of people from India after 1947 (a similar pattern does not hold for Bangladesh, as only a small number of people migrated from India). Additionally, the runs of homozygosity among Pakistani populations indicated lots of consanguineous marriages. While some South Indians marry cousins, the practice is very rare among North Indian Hindus. Rather, the genetic homogeneity of North Indian Hindus is due to the very high endogamy rates. They do not marry outside of their caste.

The results from the British Pakistanis are roughly in line with the 1000 Genome Pakistanis, but in this case, the researchers had much more granular ethnic data, as well as information on whether individuals were or were not the product of cousin-marriages. In terms of worldwide population affinity, there isn’t a great surprise. The Pathans, who are Iranian speaking, were distinct. The groups with putative Arab ancestry (Syeds), did not seem to have much of that (really, any).

The figure above shows the long-term effective population size patterns. Within the preprint the authors note that these northwest Indian populations began to diverge ~2,000 years ago. That is roughly in line with what Moorjani et al. found for their Indian samples. This tells us that these Pakistani populations were part of the same cultural milieu as Hindu populations in India itself, whose caste endogamy did not seem to crystallized until about that time. This also seems to run against the thesis presented by some Pakistani nationalists that the northwestern populations were very distinctive “non-Hindu” mlecchas. Al-Biruni and earlier observers identified caste as distinctively Indian, and the likelihood of population structure emerging at the same period in the northwest indicates that these people are broadly part of that milieu.

But I want to focus on the more recent period. Using various methods the authors estimate that the effective population sizes of many of these groups dropped 10-20 generations ago. If you assume 10 generations with generation times of 15 years, that’s 150 years. If you assume 20 generations with generation times of 25 years, that gives you 500 years. So let’s take that as our interval. What’s going on here? I think what this may illustrate is the spread of Muslim practices among Islamicized peoples of the northwest.

In my podcast with Henrich he mentions that Islamic societies are peculiar in their ubiquitous practice of “parallel-cousin-marriage.” This means that brothers will marry their children off to each other (a contrast with “cross-cousin-marriage”, common in South India, where brothers and sisters marry their children to each other). The ubiquity of cousin-marriage among Pakistani Muslims is a contrast with genetically and culturally similar populations across the border in India (Indian Punjabis do not marry cousins if Sikh or Hindu).

Click to enlarge

The fact that this practice occurred among an endogamous group for many generations has consequences. The figure to the right illustrates just how homogeneous some of these groups are against a generic European reference population. And, the fact that even unrelated individuals from the same biraderi group are often quite related. As you can see even people whose parents are unrelated still exhibit excess runs of homozygosity. This is simply a function of pedigrees being narrow, as just in Indian castes these individuals share many not-so-recent-ancestors.

A positive note is that this high level of inbreeding does not apply to Pakistani Britons where both parents were born in the country. That means that biraderi dynamics are maintained due to continuous migration from Pakistan. They’re not perpetuating themselves in the UK.

I started this post with Joe Henrich for a reason: if Henrich is correct that the differences in social structure and relatedness matter for development and economists, then Pakistan and Bangladesh might have different trajectories. Bangladesh is a corrupt and familialist society, just like Pakistan. But, that familialism is not as robust and articulated as is the norm in Pakistan. A transition to a more high-trust and non-familial society is more viable and an easier lift for a non-tribal culture where clans do not extend much beyond first cousins.

August 30, 2020

20th century genetics as basic science and 21st century genetics as basic and applied

Filed under: Genetics — Razib Khan @ 7:12 pm

There was an offhand comment on Twitter that in the 1970s genetics was barely a field because we’ve made so much progress since then. For obvious reasons, many scientists took umbrage at this. I think it’s wrong and gives the lay public the incorrect impression. But, the reality is that I do think that the way the media and some geneticists have presented the development of the field since the understanding of DNA as the substrate of inheritance in the 1950s and the explosion of genomics in the 2000s has fed into this misimpression.

What’s the truth? Genetics predates genomics by a century or more, and DNA by decades. The basics of the field were elucidated by Gregor Mendel in the 1860s. He originated the “laws of inheritance”, though unfortunately his work was ignored by contemporaries. By “laws of inheritance” I mean that Mendel formulated an analytic model that allowed for discrete inheritance and predictions of the outcome of that inheritance. Naive human understanding of heritability usually relies on an intuitive “blended theory”. It works, after a fashion, but it does not explain many patterns we see around (e.g., recessive expression).

Charles Darwin famously relied upon blended inheritance (in part) as a basis for the heritability which was essential to his theory of natural selection. But, a major problem with blended inheritance is that blending removes variation as everyone becomes a similar “mix”. This is not an issue with Mendelian inheritance, which is discrete. Alleles do not “mix”, but reconfigure every generation. Variation is retained. The “math” of evolution “works” in this manner.

The utility of Mendelian genetics is why the field exploded in the first two decades of the 20th century. Read A. H. Sturdevant’s 1913 paper on the first genetic map. I think it gives you a flavor of the rate of advancement. Genetics was definitely a field. In The Origins of Theoretical Population Genetics Will Provine outlines how this particular field of genetics developed between 1920 and 1940 to become the core of evolutionary biology. Again, this suggests that even before DNA genetics was an important field.

But, I do think it is fair to say that before 1950 genetics was very much “basic” science, and remained mostly so to the last decades of the 20th century.* DNA was interesting because it opened up the molecular biology revolution, but that had a very long fuse in terms of applications. PCR made it easier to do DNA testing, while new computing technologies made it much easier to generate and analyze data.

No one needs to be told about how genomics revolutionized the field. But it’s major impact has been transforming an often theoretical field into a massively empirical one. Modern genomics is still underpinned by the logic of Mendelian genetics. Analysis.

* The main exception here I’m going to make is for agricultural genetics, but much of this work doesn’t need “genes” as such.

August 24, 2020

Who do the English think they are?

Filed under: Anglo-Saxons,Fall of Rome,Genetics,History,Population genetics — Razib Khan @ 3:04 pm

In the early 5th century the Roman legions abandoned Britain, and the sceptered isle fell off the pages of history. When it reemerges two centuries later Celtic Britain had become the seedbed for the nation-state of England. The Christian religion, newly-established on the island at the time, had given way once again to paganism. Brythonic Celtic speech was ascendant only on the fringes. A cacophony of German dialects spread out across the fertile south and east, radiating out of the “Saxon Shore”.

This ethno-religious transformation of the island occurred under the shadow of semi-history, allowing for the development of an imaginative romantic tradition exemplified by the Arthurian Cycle. But this Dark Age also became a bone of contention between the English who saw themselves as deeply rooted in the land, and those who declared that they were a German folk who had won their new home through conquest and blood. The dominant view at any given time reflected social and political events of the 20th century more than facts. The propaganda value of myth meant more than the conjectures of scholars.

While in the early 20th century the dominant position was that the English were a people akin to German Saxons, a race apart from the Welsh, by the early 21st century serious scholars assumed that the spread of Anglo-Saxon culture occurred through imitation rather than replacement.

Today we can say with some confidence that neither stark view is correct, and that the middle path between is far more interesting and complex. Large numbers of Saxons, Angles and Jutes did in fact cross the North Sea — but the preponderance of England’s heritage still draws from the Celtic-speaking peoples. It is not coincidence that the earliest rulers in Alfred the Great’s lineage bear Celtic names, not German ones.

Those who argued for the erasure of the Celtic people did not do so without any basis. St. Gildas, a 6th century British Celt, recounted in On the Ruin and Conquest of Britain the defeat and destruction of his people at the hands of the Saxons. More recently, 19th century philologists observed that the number of Brythonic Celtic loan words in English is extremely small; in fact, there may be more Celtic loan words from Gaulish, due to the later Norman French influence. Finally, the collapse of institutions like the Roman Christian Church and the total decay of urban life indicates incredible disruption of the social hierarchy which characterised post-Roman Britain.

A contrast here exists with Gaul, which absorbed a German-speaking elite but retained Roman language and religion. Some of the nobility of southern and western France even traced their descent from Romans, not Germans. On a more demotic level, British archaeologists have also observed that the arrival of the Saxons seems to have been associated with a transformation of the layout of rural farmsteads. In most societies, farmers have customs and traditions which they hew to, and they are often quite stubborn and set in their ways. Such a change indicates new people, not just practices.

But by the late 20th century such views of cultural and demographic disruption were in bad odour. The dominant ethos is that people did not move, their customs and traditions did. Hengist and Horsa may have existed, but rather than a folk migration the Anglo-Saxon conquest was one of a small number of German mercenaries who were engaged in elite capture of the post-Roman peasantry.

The Welsh historian Norman Davies observed in his 1999 book The Isles that “blood price” in 8th century Wessex differed between whether one was Saxon or British, the implication here being that there were many Celtic Britons living in the Anglo-Saxon lands, even if our documentary evidence is from the Saxon elite; this would tally with the 6th century ancestors of Alfred the Great having names such as Ceawlin, Cynegils and Cerdic, all of which have a distinctive Welsh flavor.

Genetics has untangled the Gordian knot of this semi-historical mystery, although illumination has not come at once, and only in fits and starts. One of the primary reasons is that the genetic difference between “Celtic” and “German” peoples is very small. Most Northern Europeans separated from each other very recently. Ancient DNA from between three and eight thousand years ago shows that Northern Europe underwent several mass migrations which transformed the genetic landscape.

First, the blue-eyed dark-skinned hunter-gatherers who descend from Ice Age Europeans disappeared and were absorbed by brown-eyed pale-skinned farmers who moved north out of the Near East. Then, these agriculturists were themselves overwhelmed by a people who migrated out of the Eurasian steppe into Europe 5,000 years ago. These pastoralist people probably brought Indo-European languages, and 4,500 years ago they arrived in Britain as the Bell Beaker culture. Within a few generations there was 90% genetic turnover, as the farmers who first erected Stonehenge disappeared, and were replaced by people who seem to have arrived from what is today northern Germany, possibly prefiguring the later Anglo-Saxon migration.

The problem from the perspective of genetics in understanding the proportion of Anglo-Saxon ancestry in the modern English goes back to the reality that Germans and Celts themselves had only been separated for 3,000 years, at most. These are genetically very close populations, and the technology of the early 21st century could not resolve the questions being asked.

UCL geneticist Steve Jones did attempt such a thing in his 2003 book Y: The Descent of Men. Jones observed that the distribution of two Y chromosomal lineages exhibits a sharp break at Offa’s Dyke. A far higher proportion of Welsh men are R1b, which is very common across the Atlantic facade of Europe, while more English men carry R1a, which is found in higher frequencies in Germany and Norway. In contrast, Professor Jones observed that there was no difference in the maternal heritage of the Welsh and English, suggesting that the ethnic change was due to the impact of men. Jones’s UCL colleague Mark Thomas later developed an “apartheid model” to explain why the genetic difference between the English and Welsh was so striking.

But the true understanding of the situation could only be obtained by looking across the whole genome, not simply the paternal and maternal lineages. This was done by the Peopling of the British Isles Project, which published a paper in 2015 that drew from analysis on hundreds of thousands of genetic markers from 2,000 British individuals who were sampled from all across the United Kingdom.

They estimated that 10-40% of the ancestry in central and southern England was Anglo-Saxon — that is, DNA segments more similar to the Germans than the Welsh. Another paper from 2016, utilising ancient as well as contemporary DNA, estimated that 38% of the ancestry in the “East English” — people from East Anglia and the East Midlands — is derived from the Anglo-Saxons. These researchers actually found DNA from Dark Age-era graves identified as Anglo-Saxon, and some of these individuals were far more like the Germans in their DNA than the modern English; they differed from earlier Iron Age samples, proving beyond a doubt that a significant number of Germans did cross the North Sea in the 6th century.

Where does this leave us in relation to the question of whether the transformation of Dark Age Britain to early medieval England was one of genes or memes? The clear answer seems to be both. The emergence of a new style of farming, pottery and the collapse of urban Roman civilization and Christianity in eastern Britain was not simply due to the prestige and power of a small number of German warlords. Whole villages must have transplanted themselves across the North Sea, creating the nucleus of a new people, and absorbed the remaining British Celts. The lack of Celtic loanwords and the adoption of Saxon peasant culture may indicate the self-confidence of the newcomers. If St. Gildas is correct, the British elites moved to the west of the island, leaving the common people to their own devices.

But though the southern and eastern fringe of England has a substantial Anglo-Saxon demographic imprint, that fades out as one moves to the west, including to the lands that once comprised the kingdom of Wessex. There is far less German genetic influence in Hampshire, Berkshire or Wiltshire, let alone Devon. We know from early medieval records that Celtic language speakers did exist as late as the 8th century in these domains (and much later in Devon) but by then Old English, which is for all purposes a purely Germanic language, was dominant.

The genealogy of the House of Wessex may offer a clue as to what occurred in broad swaths of western England. In the 6th century Celtic names imply that this elite lineage was identified with British culture, and looked west, but by the 7th German names became common, and the kings were pagan. Though the Saxons may have imposed their way of life through sheer numbers in the east, explaining the light impact of British Celtic culture upon their folkways and language, their expansion beyond the Saxon Shore seems to have been due to the adoption of the German identity by native British. The killing of a Celtic-speaking individual under the Saxon system of blood price was far cheaper than for a German speaker, serving as a clear inducement to assimilate.

What science makes clear then is that both extreme scenarios presented in the 19th and 20th centuries were wrong. The English are not a race apart from the Welsh. The modern English are genetically closest to the Celtic peoples of the British Isles, but the modern English are not simply Celts who speak a German language. A large number of Germans migrated to Britain in the 6th century, and there are parts of England where nearly half the ancestry is Germanic.

These folk served as the focus of a cultural revolution that transformed the British Isles. It was not a passive affair: the cities, churches, and hamlets of the previous inhabitants were blotted out, and what had been one of the provinces of the Roman Empire became a backwater pagan land. Though the original Romano-British elites had some knowledge of Latin, and patronised the Christian Church, the patina of civilization was clearly thin upon them, and the loosely Christian Celtic warlords of Dark Age western Britain transformed seamlessly into the pagan kings of Anglo-Saxon England.

The initial founding of the Saxon Shore was surely based on a level of brutality that Christian priests, if any had lived to tell the tale, would have recorded with foreboding. But the transformation of vast swaths of western Britain into the core of what had become England by the Viking Age occurred consensually, so seductive had the Saxon society become to the Celts, highborn and low.

The lesson that history and genetics teach us that cultural change is a complex phenomenon, and a single factor does not explain the whole story. Today we live in an age of migration, and native peoples fear being replaced, while immigrant communities fear being assimilated. Numbers matter, but the Saxons tell us that numbers are not everything.

Who do the English think they are?

Filed under: Anglo-Saxons,Fall of Rome,Genetics,History,Population genetics — Razib Khan @ 3:04 pm

In the early 5th century the Roman legions abandoned Britain, and the sceptered isle fell off the pages of history. When it reemerges two centuries later Celtic Britain had become the seedbed for the nation-state of England. The Christian religion, newly-established on the island at the time, had given way once again to paganism. Brythonic Celtic speech was ascendant only on the fringes. A cacophony of German dialects spread out across the fertile south and east, radiating out of the “Saxon Shore”.

This ethno-religious transformation of the island occurred under the shadow of semi-history, allowing for the development of an imaginative romantic tradition exemplified by the Arthurian Cycle. But this Dark Age also became a bone of contention between the English who saw themselves as deeply rooted in the land, and those who declared that they were a German folk who had won their new home through conquest and blood. The dominant view at any given time reflected social and political events of the 20th century more than facts. The propaganda value of myth meant more than the conjectures of scholars.

While in the early 20th century the dominant position was that the English were a people akin to German Saxons, a race apart from the Welsh, by the early 21st century serious scholars assumed that the spread of Anglo-Saxon culture occurred through imitation rather than replacement.

Today we can say with some confidence that neither stark view is correct, and that the middle path between is far more interesting and complex. Large numbers of Saxons, Angles and Jutes did in fact cross the North Sea — but the preponderance of England’s heritage still draws from the Celtic-speaking peoples. It is not coincidence that the earliest rulers in Alfred the Great’s lineage bear Celtic names, not German ones.

Those who argued for the erasure of the Celtic people did not do so without any basis. St. Gildas, a 6th century British Celt, recounted in On the Ruin and Conquest of Britain the defeat and destruction of his people at the hands of the Saxons. More recently, 19th century philologists observed that the number of Brythonic Celtic loan words in English is extremely small; in fact, there may be more Celtic loan words from Gaulish, due to the later Norman French influence. Finally, the collapse of institutions like the Roman Christian Church and the total decay of urban life indicates incredible disruption of the social hierarchy which characterised post-Roman Britain.

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A contrast here exists with Gaul, which absorbed a German-speaking elite but retained Roman language and religion. Some of the nobility of southern and western France even traced their descent from Romans, not Germans. On a more demotic level, British archaeologists have also observed that the arrival of the Saxons seems to have been associated with a transformation of the layout of rural farmsteads. In most societies, farmers have customs and traditions which they hew to, and they are often quite stubborn and set in their ways. Such a change indicates new people, not just practices.

But by the late 20th century such views of cultural and demographic disruption were in bad odour. The dominant ethos is that people did not move, their customs and traditions did. Hengist and Horsa may have existed, but rather than a folk migration the Anglo-Saxon conquest was one of a small number of German mercenaries who were engaged in elite capture of the post-Roman peasantry.

The Welsh historian Norman Davies observed in his 1999 book The Isles that “blood price” in 8th century Wessex differed between whether one was Saxon or British, the implication here being that there were many Celtic Britons living in the Anglo-Saxon lands, even if our documentary evidence is from the Saxon elite; this would tally with the 6th century ancestors of Alfred the Great having names such as Ceawlin, Cynegils and Cerdic, all of which have a distinctive Welsh flavor.

Genetics has untangled the Gordian knot of this semi-historical mystery, although illumination has not come at once, and only in fits and starts. One of the primary reasons is that the genetic difference between “Celtic” and “German” peoples is very small. Most Northern Europeans separated from each other very recently. Ancient DNA from between three and eight thousand years ago shows that Northern Europe underwent several mass migrations which transformed the genetic landscape.

First, the blue-eyed dark-skinned hunter-gatherers who descend from Ice Age Europeans disappeared and were absorbed by brown-eyed pale-skinned farmers who moved north out of the Near East. Then, these agriculturists were themselves overwhelmed by a people who migrated out of the Eurasian steppe into Europe 5,000 years ago. These pastoralist people probably brought Indo-European languages, and 4,500 years ago they arrived in Britain as the Bell Beaker culture. Within a few generations there was 90% genetic turnover, as the farmers who first erected Stonehenge disappeared, and were replaced by people who seem to have arrived from what is today northern Germany, possibly prefiguring the later Anglo-Saxon migration.

The problem from the perspective of genetics in understanding the proportion of Anglo-Saxon ancestry in the modern English goes back to the reality that Germans and Celts themselves had only been separated for 3,000 years, at most. These are genetically very close populations, and the technology of the early 21st century could not resolve the questions being asked.

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Britain's divisions go way, way back

By Ed West

UCL geneticist Steve Jones did attempt such a thing in his 2003 book Y: The Descent of Men. Jones observed that the distribution of two Y chromosomal lineages exhibits a sharp break at Offa’s Dyke. A far higher proportion of Welsh men are R1b, which is very common across the Atlantic facade of Europe, while more English men carry R1a, which is found in higher frequencies in Germany and Norway. In contrast, Professor Jones observed that there was no difference in the maternal heritage of the Welsh and English, suggesting that the ethnic change was due to the impact of men. Jones’s UCL colleague Mark Thomas later developed an “apartheid model” to explain why the genetic difference between the English and Welsh was so striking.

But the true understanding of the situation could only be obtained by looking across the whole genome, not simply the paternal and maternal lineages. This was done by the Peopling of the British Isles Project, which published a paper in 2015 that drew from analysis on hundreds of thousands of genetic markers from 2,000 British individuals who were sampled from all across the United Kingdom.

They estimated that 10-40% of the ancestry in central and southern England was Anglo-Saxon — that is, DNA segments more similar to the Germans than the Welsh. Another paper from 2016, utilising ancient as well as contemporary DNA, estimated that 38% of the ancestry in the “East English” — people from East Anglia and the East Midlands — is derived from the Anglo-Saxons. These researchers actually found DNA from Dark Age-era graves identified as Anglo-Saxon, and some of these individuals were far more like the Germans in their DNA than the modern English; they differed from earlier Iron Age samples, proving beyond a doubt that a significant number of Germans did cross the North Sea in the 6th century.

Where does this leave us in relation to the question of whether the transformation of Dark Age Britain to early medieval England was one of genes or memes? The clear answer seems to be both. The emergence of a new style of farming, pottery and the collapse of urban Roman civilization and Christianity in eastern Britain was not simply due to the prestige and power of a small number of German warlords. Whole villages must have transplanted themselves across the North Sea, creating the nucleus of a new people, and absorbed the remaining British Celts. The lack of Celtic loanwords and the adoption of Saxon peasant culture may indicate the self-confidence of the newcomers. If St. Gildas is correct, the British elites moved to the west of the island, leaving the common people to their own devices.

But though the southern and eastern fringe of England has a substantial Anglo-Saxon demographic imprint, that fades out as one moves to the west, including to the lands that once comprised the kingdom of Wessex. There is far less German genetic influence in Hampshire, Berkshire or Wiltshire, let alone Devon. We know from early medieval records that Celtic language speakers did exist as late as the 8th century in these domains (and much later in Devon) but by then Old English, which is for all purposes a purely Germanic language, was dominant.

Suggested reading
It wasn't the Berlin Wall that divided Germany

By James Hawes

The genealogy of the House of Wessex may offer a clue as to what occurred in broad swaths of western England. In the 6th century Celtic names imply that this elite lineage was identified with British culture, and looked west, but by the 7th German names became common, and the kings were pagan. Though the Saxons may have imposed their way of life through sheer numbers in the east, explaining the light impact of British Celtic culture upon their folkways and language, their expansion beyond the Saxon Shore seems to have been due to the adoption of the German identity by native British. The killing of a Celtic-speaking individual under the Saxon system of blood price was far cheaper than for a German speaker, serving as a clear inducement to assimilate.

What science makes clear then is that both extreme scenarios presented in the 19th and 20th centuries were wrong. The English are not a race apart from the Welsh. The modern English are genetically closest to the Celtic peoples of the British Isles, but the modern English are not simply Celts who speak a German language. A large number of Germans migrated to Britain in the 6th century, and there are parts of England where nearly half the ancestry is Germanic.

These folk served as the focus of a cultural revolution that transformed the British Isles. It was not a passive affair: the cities, churches, and hamlets of the previous inhabitants were blotted out, and what had been one of the provinces of the Roman Empire became a backwater pagan land. Though the original Romano-British elites had some knowledge of Latin, and patronised the Christian Church, the patina of civilization was clearly thin upon them, and the loosely Christian Celtic warlords of Dark Age western Britain transformed seamlessly into the pagan kings of Anglo-Saxon England.

The initial founding of the Saxon Shore was surely based on a level of brutality that Christian priests, if any had lived to tell the tale, would have recorded with foreboding. But the transformation of vast swaths of western Britain into the core of what had become England by the Viking Age occurred consensually, so seductive had the Saxon society become to the Celts, highborn and low.

The lesson that history and genetics teach us that cultural change is a complex phenomenon, and a single factor does not explain the whole story. Today we live in an age of migration, and native peoples fear being replaced, while immigrant communities fear being assimilated. Numbers matter, but the Saxons tell us that numbers are not everything.

The post Who do the English think they are? appeared first on UnHerd.

August 10, 2020

Kashmiri Brahmins are just like other Kashmiris

Filed under: Genetics — Razib Khan @ 3:28 pm

I think I’ve posted this before, but it was a while ago before we had so many readers. In this paper they took 15 random Kashmiris from the Valley, and compared them to various populations. The plot below, as well as admixture analysis in the paper, shows no daylight between the Pandit samples and generic Muslim Kashmiris.

This is not to say Pandits are not an endogamous community and were not before the Islamicization of Kashmir. But, it is to say that in their overall genome their origins are exactly the same as other Kashmiris. This is in contrast to many parts of India in regards to Brahmins, though the “stylized fact” seems to be the further north and west you go, the smaller the genome-wide difference between Brahmins and non-Brahmins will be. This seems to comport with the idea that Brahmins are intrusive to the south and east in a way they are not to the north and west.

Finally, the data from ancient DNA is strongly suggestive of “AASI-reflux” across north and west South Asia after 3000 BC. See my post The Aryan Integration Theory (AIT).

July 23, 2020

What do we call the Ancient Ancestral North Indians?

Filed under: Genetics — Razib Khan @ 12:06 pm

Commenters on this weblog have expressed dissatisfaction with the nomenclature of the “eastern Iranian farmers” who were the dominant genetic contributors to the Indus Valley People. The author of The formation of human populations
in South and Central Asia
agrees that this is a problem.

To review: the dominant ancestry component, called Iranian-related or eastern Iranian farmer, has two components. About 5-10% is related to “West Siberian Hunter-Gatherers”, who mostly descend from “Ancient North Eurasian” Paleo-Siberian groups (this group contributed ancestry to eastern European hunter-gatherers and Native Americans). The remainder of the ancestry is related to farming populations that are termed “Iranian” from samples in the Zagros in the early Holocene. But the genetics indicates that the separation of the Indian ancestry component dates to before farming, probably between 10-15,000 years ago. Without ancient DNA that is older, we can’t be sure of its geographic range, but it is reasonable to infer that this was an eastern expansion of hunter-gatherers out of the Zagros (seeing as how the WSHG ancestry is not found in the west, and the broader Iranian farmer clade seems to form a clade with Anatolian farmers and Levantine farmers).

But obviously the use of the term “Iranian” confuses with the nation-state of Iran.  This has come up when I use terms like “Iranian-speaking people,” and people get confused because they don’t assume that I’m talking about people who live in Russia (Ossetes), or ancient people who flourished in Xinjiang and Ukraine.

Historically modern Iran was called “Persia”, and Iran was actually more of an archaic civilizational term. But in the 20th-century the Pahlavi’s resurrected this ancient term for the nation-state, so here we are.

The question this: what is a better term for the “Iranian-related farmers”? I have often used the awkward “NW South Asia”, since it seems plausible this group was present in modern-day Pakistan by the early Holocene, and probably earlier. Thoughts?

I’m basically asking for terms and why you think those terms are good. I may adopt a term in the comments for usage on my blogs.

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