As many of you know when you have two adjacent demes, breeding populations, they often rapidly equilibrate in gene frequencies if they were originally distinct. There are plenty of good concrete examples of this. The Hui of China are Muslims who speak local Chinese dialects. The most probable root of this community goes back to the enormous population of Central Asia Muslims brought by the Mongol Yuan dynasty that ruled ruled China for over a century from the late 1200s to 1300s. Genetic studies of this group that I’ve seen indicate that a high bound estimate for West Eurasian ancestry is ~10%. The other ~90% is interchangeable with the Han Chinese. So let’s assume that the Hui are ~10% West Asian. If you assume that in the year 1400 the Hui were “pure,” you have 24 generations (25 years per generation). The original population of “Central Asian Muslims” were heterogeneous, including Iranians and Turks. But let’s take it granted that they were 50% East Eurasian and 50% West Eurasian in ancestry at the time of their arrival. What would the intermarriage rate per generation have to be so that the Hui are ~10% West Eurasian at t = 24 (24 generations after the beginning of intermarriage assuming 50/50 West vs. East Eurasian splits)? Turns out all you need is a constant 7% intermarriage rate per generation (the Han Chinese population is so large in relation to the Hui that you can model it as infinite in size).
The situation gets even simpler when you have one population which divides into two. For example, imagine that the Serbs and Croats fissioned from a set of unstructured South Slavic tribes which filtered into ancient Illyria ~600 A.D. Soon enough there was a cultural division between the two in terms of religion (Western vs. Eastern Christian) which threw up a population genetic barrier. If you assume that genetically the two groups were totally similar at t = 0, and you separated them perfectly, over time they would diverge due to drift in their allele frequencies. But the reality is that barriers between geographically close groups do not prevent all intermarriage. Even extremely insular groups in a cultural sense such as the Roma of Eastern Europe are clearly heavily admixed with their surrounding populations, as they seem to be no more than ~50% South Asian in total genome content. Going back to the South Slavs, who start out very similar in our putative scenario, how much intermarriage will be necessary for them to not diverge? The issue is not the rate of intermarriage, rather, one migrant per generation across the two demes will be sufficient to equilibrate allele frequencies. On the face of it this seems implausible, but recall that divergence is driven mostly by drifting of genes as well as new variation (whether through other exogenous migratory sources or mutation). Very small populations are subject to a lot of drift, and so diverge rapidly, but only very few migrants are needed to bring it back into alignment, because they are proportionally significant. In contrast, the frequencies of large populations are less buffeted by generation-to-generation sample variance (e.g., 10 tosses of a coin will deviate more from 50/50 proportionally than 100 tosses), requiring less gene flow proportionally to maintain parity.
Dienekes Pontikos keeps chugging along, and has cranked out a new bar plot from the ADMIXTURE program with 15 putative ancestral components. He has “69 populations, and 1,189 individuals in total.” Most of these were assembled from public data, but some of them are particular to the Dodecad Ancestry Project. He contends:
In comparison to the K=10 analysis, the increased resolution allows us to:
– South Asians belonged primarily to the South Asian and West Asian components; this South Asian component spilt over to Iran and Central Asia. Now, a new Central-South Asian component, corresponding to the Ancestral North Indian of a recent study is inferred, and a corresponding South Indian component.
– HGDP Bedouins and Behar et al. (2010) Saudis take up their own component which I labeled Arabian. This appears to be a subset of the Southwest Asian component of the K=10 analysis
– There are several components in Siberian and Central Asian populations, alread discovered in my regional analysis. These are Central Siberian, Nganasan, Koryak, Chukchi, and Altaic which replace the K=10 Northeast Asian component
Not only has he generated a bar plot, but there is a PCA showing the relationship between the 15 ancestral groups, as well as a hierarchical tree. Since he references to the ANI and ASI of Reich et al., I thought I would note that the South Indian element from Dienekes’ K = 15 is still found in appreciable portions in the Turkic groups which earlier exhibited the South Asian component. And, on the PCA and phylogenetic tree it still clusters with West Eurasians more than East Eurasians, which is not the case with ASI (or the various Indian mtDNA lineages which coalesce back to a more recent common ancestor with East Eurasians).
The bar plot is below. Of interest are the most “pure” European groups, the Sardinians and Lithuanians. Also, compare Scandinavians and Finns.
If you haven’t, you should keep an eye on Dienekes‘ Dodecad Ancestry Project (RSS). The pilot phase of data collection is over, and the first population level statistics are now coming out. Of particular interest to me is a new analysis of various northern European ethnicities just published.
The samples used in this analysis are:
– 25 HapMap-3 White Americans. These are the Mormons of predominantly Northwest European heritage
– 5 Dodecad Project Finns
– 25 HGDP-CEPH Russians from Vologda, in north-central European Russia
– 12 Dodecad Project continental Germanics (Scandinavians and Germans)
– 10 Behar et al. (2010) Lithuanians
– 9 Behar et al. (2010) Belorussians
– 3 Dodecad Project Northern Slavs
Below are two visualizations of the genetic structure. First, an MDS. And second, a bar plot of ancestral quanta derived from ADMIXTURE. I’ve added some clarifying labels.
If you live in the States one of the things you hear a lot about Europe in regards to its relationship to its ethno-religious minorities are the problems with Muslims. This is probably an Americo-centric perspective shaped by 9/11, when many of the hijackers had turned out to have spent time in Germany. Additionally, terrorist actions in both London and Madrid highlight the persistence of these problems over the years. These sorts of shocking events put a sharp focus on the geopolitical cross-hairs which Europe finds itself in in the second age of mass migration. Though this time it is a destination, and not a source.
But having been to Europe recently it was notable that in several regions the day-to-day tension when it came to ethnicity often focused on Gypsies (I use the older term because the ethnonym “Roma” which has become politically correct in the USA includes only a subset of Europe’s Gypsy population, even if the greater number). Many regions of Europe now have two distinct populations of Gypsies, a long resident local group, as well as Roma from the eastern nations of the EU. Though the relationships between these traditionally nomadic peoples and indigenous populations has never been without tension, it is clear that something close to a modus vivendi has been achieved in many European nations between the majority and their small native Gypsy populations. The influx of the Balkan Roma add a new variable.
But the political fuss for me simply rekindled a curiosity as to the genetic origins of the Gypsies. Culturally their South Asian provenance couldn’t be clearer; they speak an Indo-Aryan language. Their term for themselves in many parts of Europe comes from the Indo-Aryan word for “black,” as they are are darker than the natives of the lands in which they have settled , and in fact often look visibly South Asian. This seemed especially true of Balkan Roma. On the other hand the Kale of Finland looked to be brunette Europeans.
Sometimes in applied fields artistic license is constrained by the necessity of function to particular creative channels. Architecture comes to mind, at least before innovative technologies produced lighter and stronger materials, freeing up form from its straitjacket (whether this was a positive development is a matter of taste). But there’s only so much you can do with your palette when your palette is limited. This can be a bug, or it can be a feature. Science is not art, but in some ways at its heart it’s a story about the universe. The story can be in words or math, no matter, ultimately it’s the human attempt to map nature and make its subtle patterns comprehensible to us in plainer fashion. Some of the human biases in our quest are transparent. Why is there anthropology? A whole discipline devoted to the study of mankind and his nearest biological kin. We don’t peruse the patters with an objective and uninterested eye. We’re shaped by our presuppositions, as well as the constraints of the methods, and the results we have before us. The emergence of a theoretical evolutionary biology in the decades before the molecular revolution after World War II may have been in part simply a function of the fact that there were only so many results one could squeeze out of classical evolutionary genetic techniques, which relied on tracking only a limited set of phenotypes due to large effect mutations in breeding populations. With the rise of molecular evolution you saw the crystallization of theoretical frameworks, such as the neutral theory, to explain the burst of novel results.
Around the year 2000 something similar happened in historical population genetics. The analysis of mtDNA lineages, passed from mother to daughter, had matured, and techniques for typing the Y chromosome had started to catch up, so that a symmetry between the sexes could arise. “Mitochondrial Eve” was now paired with “Y chromosomal Adam.” Though mtDNA and Y lineages were only two direct lines of ancestry, because there was no recombination across much of their sequence it was easy to analyze them within the context of coalescent theory. In contrast, the genealogy of autosomal regions of the genome were confounded by recombination, which mixed & matched the variation in a manner which made reconstruction of past history far more difficult. So we had the technology to extract the genetic variation from mtDNA and the Y chromosome, and, we knew how to model their evolution. The two together produced a genetic time machine.
In my post on Empires of the Word I observed that quite often the written record is silent on many matters which only language or genes tell us must have occurred. The Indo-Aryan character of the dominant language on the island of Sri Lanka seems to be a geographical anomaly in the least, but perhaps most strange of all is the existence of a language and ethnic group of clear Southeast Asian provenance on the island of Madagascar. To my knowledge Arab, Persian and South Asian sources do not record the existence of a prominent Southeast Asian maritime diaspora which spanned the Indian ocean in the years before 1000 A.D., but we know that it did exist. A new paper on the genetics of the island of Comoros fleshes out another piece of the puzzle, Genetic diversity on the Comoros Islands shows early seafaring as major determinant of human biocultural evolution in the Western Indian Ocean:
I am currently reading Peter Heather’s Empires and Barbarians: The Fall of Rome and the Birth of Europe. This is a substantially more hefty volume in terms of density than The Fall of the Roman Empire: A New History of Rome and the Barbarians . It is also somewhat of a page turner. One aspect of Heather’s argument so far is his attempt to navigate a path between the historically tinged fantasy of what its critics label the “Grand Narrative” of mass migration of barbarian tribes such as the Goths, Vandals and Saxons during the 4th to 6th centuries, dominant before World War II, and its post-World World II counterpoint. As a reaction against this idea archaeologists have taken to a model of pots-not-people, whereby cultural forms flow between populations, and identities are fluid and often created de novo. This model would suggest that only a tiny core cadre of “German” “barbarians” (and yes, often in this area of scholarship the most banal terms are problematized and placed in quotations!) entered the Roman Empire, and the development of a Frankish ruling class in the former Gaul, for example, was a process whereby Romans assimilated to the Germanic identity (with the shift from togas to trousers being the most widespread obvious illustration of Germanization of norms). I believe that liberally applied this model is fantasy as well. Being a weblog where genetics is important, my skepticism of both extreme scenarios is rooted in new scientific data.
Jared Diamond famously argued in Guns, Germs and Steel that only a small set of organisms have the characteristics which make them viable domesticates. Diamond’s thesis is that the distribution of these organisms congenial to a mutualistic relationship with man shaped the arc of our species’ history and the variation in wealth that we see (though his a human-centric tale, we may enslave them, eat and use them as beasts of burden, but these are also species which have spread across the world with our expansion). This thesis has been challenged, but the bigger point of putting a focus on how humans relate to their domesticated animals, and the complex co-evolutionary path between the two, is something that we need to consider. In a plain biological and physical sense animals have utility; we eat them, and for thousands of years they were critical to our transportation networks. Some have argued that the rise of Islam, Arab monotheism, was contingent on the domestication of the camel (which opened up interior trade networks previously unaccessible). In The Horse, the Wheel, and Language: How Bronze-Age Riders from the Eurasian Steppes Shaped the Modern World the argument is made that the distribution of the Indo-European languages has to do with the facility of Central Eurasian plainsmen with their steeds. And of course there is the domestic dog, arguably the one creature which is able to read our emotions as if they were a con-specific.
I suspect that the evolution and ethology of domesticated animals will offer a window into our own evolution and ethology. Konrad Lorenz famously believed that humans were going through their own process of domestication all the while that they were selecting organisms suited to their own needs. More pliable, less intelligent, faster growing and maturing, and so forth. Know thy companions, and know thyself, so to speak.
What about an animal as intelligent as a dog, but famously tasty? (the combination of the two characters causing some ethical tension in the minds of many) I speak here of the pig. A few years ago research came out which showed that pig-culture was introduced to Europe from the Middle East. That is, Middle Eastern pigs came with Middle Eastern people in all likelihood. But modern European pigs do not derive from these lineages, rather, by comparing modern genetic variation with ancient DNA the authors showed that the Neolithic pigs had been replaced by local breeds. Just as pigs can go feral and fend for themselves rather easily, it seems that their basic morph can be derived from wild boar populations easily as well (by contrast, it will perhaps take some effort to derive a pekingese from wolf populations, offering a reason for why small dogs seem to have emerged once). A new paper explores the evolutionary history and phylogeography of the pigs of the swine-loving societies par excellence, those of East Asia. Patterns of East Asian pig domestication, migration, and turnover revealed by modern and ancient DNA:
The following passage is from the epilogue of The Real Eve: Modern Man’s Journey Out of Africa by Stephen Oppenheimer:
In this book I have offered a synthesis of genetic and other evidence. Everything points to a single southern exodus from Eritrea to the Yemen, and to all the non-African male and female gene lines having arisen from their respective single out-of-Africa founder lines in South Asian (or at least near the southern exit). I regard the genetic logic for this synthesis as a solid foundation, and I have based the rest of my reconstruction of the human diaspora upon it. Obviously, the ‘choice’ of starting point (mine or theirs) determined all the subsequent routes our ancestors and cousins took. Tracing the onward trails is only possible as a result of marked specificity in regional distribution of the genetic branches The geographic clarity of both male and female gene trees is a big departure from the fuzzy inter-regional picture shown by older genetic studies. The degree of segregation of lines into different countries and continents is in itself good evidence that once they got to their chosen new homes, the pioneers generally stayed put, at least until the Last Glacial maximum forced some of them to move. This conservative aspect of our genetic prehistory also provides a partial explanation for the fact that when we look at a person, we can usually tell, to the continent, where their immediate ancestors came from, and underlies differences that some of us still call ‘race.’
Oppenheimer wrote the above in the early aughts, as his book was published in 2003. Much of this is generally in line with the ‘orthodoxy’ of the day. I believe that Oppenheimer’s assertion that there was one southern migration out of Africa by anatomically modern humans has gained some advantage over the alternative model of two routes, northern and southern, over the past ten years (Spencer Wells’ The Journey of Man sketches out the two wave model). Other assertions and assumptions have not stood the test of time. In particular, I would contend that generally the ‘conservative aspect of our genetic prehistory’ can no longer be taken for granted. Specifically, it seems likely now that much occurred after the Ice Age and during the Neolithic.