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

April 1, 2018

When do people forget where they come form?

Filed under: Migration,Prehistory — Razib Khan @ 9:09 pm

When it comes to the arrival of Indo-Aryans to South Asia a major question Indians always post is “if they are invaders why don’t they mention that in their mythology?” My standard rejoinder is straightforward: we have plenty of paleogenetic evidence that many populations are intruders, but their mythology doesn’t indicate that. If the Indian objection is to hold then why not others? Are all human populations autochtonous in their native lands?

And yet the most recent work suggests that steppe ancestry didn’t arrive in the BMAC region until 2000 BC. That means that the Cemetary H culture in Punjab dating to 1900 BC is the earliest likely candidate for Indo-Aryans in South Asia. The Rigveda was composed as early as 200 years after this date, or as late as 700 years. Could they have “forgotten” where they came from?

The Irish are one people who have preserved their mythology due to the gradual and indigenous nature o Christianization. But 2,500 years after their arrival en masse from the continent they had forgotten the details. But, the motif of invasion was preserved, though we don’t know if that is a memory of their past, or just a channeling of the mythos of the period when their folklore was written down. Another example might be the Japanese, who arrived about 1,100 years before the Heian period, the first flowering of literate civilization on the island. To my knowledge, their mass migration from southern Korea was mostly forgotten by then.

With all this in mind, I decided to reread Genetic origins of the Minoans and Mycenaeans. The conclusion of the paper is that it’s clear an appreciable, though minority, component of Mycenaean ancestry seems to have some affinity with Indo-European groups. The two candidates for the donors are people from the Eurasian steppe, or Copper age Armenians. For linguistic reasons that I can barely evaluate, I lean toward the former. This implies that the proto-Greeks arrived in the late 3rd millennium or early 2nd millennium. The mythology of ancient Greek as recorded by Hesiod and others in the Archaic period probably dates in part to the Bronze Age (some of the Greek gods are recorded in the Linear B tablets). To my knowledge the Greeks do not record when they arrived from outside of Greece.

This suggests that 1,000 years is sufficient for a forgetting, at least for a semi-literate society.

The last is key. Societies with written histories can maintain continuity. But what about oral societies?

February 21, 2018

The genetics of the Lombard folk migration

Filed under: barbarians,Germans,Lombards,Migration,Roman History,Romans — Razib Khan @ 9:13 am


There are many debates about the period after the fall of the Western Roman Empire in the late 5th century. For example, did it “fall” in the first place? I believe that the concomitant p0litical, social, and economic changes do warrant that word. But another question concerns the “barbarians,” who were mostly German peoples (there are some exceptions, such as the Iranian Alans and the Huns, whose specific provenance is unclear). Were they ethnically and politically coherent? Were they even peoples?

The extreme stylized positions might be outlined as follows:

– The barbarians who filled the political vacuum after the collapse of the late Roman state were coherent preexistent ethnic and political entities of German origin who migrated en masse and engaged in a folk wandering.

– Though their original provenance may have been in bands of German warriors from specific tribes, but the time they appear on the stage of history as we understand it, the barbarians were in fact a motley crew of opportunists of various origins, who adhered to a “barbarian” identity which was created de novo with the collapse of Rome. They were made by the collapse, they did not cause the collapse.

In the late 1990s, Norman Davies in The Isles presents an argument closer to the latter for the British Isles. That is, the Anglo-Saxon character of Britain was to a large effect a function of elite emulation and diffusion of a Germanic culture introduced by what was operationally a late Roman mercenary class. Davies alludes to texts which indicate a substantial native British population in Anglo-Saxon England centuries after the fall of Celtic kingdoms. This is in contrast to the apocalyptic vision of British monk Gildas, who depicts his Brythonic people fleeing before pagan Saxons and being driven into the sea. And, I have alluded to the possibility that the West Saxon monarchy, which later came to the center of English history during the Viking incursion, was in fact in origin Romano-British, rather than German (the early kings have Celtic names).

And yet England was always the most difficult case for cultural diffusion, because to a great extent Roman-British society did collapse. Both the British Celtic language and Christianity seem to have faded from the landscape, so the that the latter had to be reintroduced by Irish and continental European missionaries. Today, the genetics is more definitive, and it seems a substantial German migration did impact what became England, especially the east, what was the Saxon Shore. Though the majority of the ancestry of the people of England today seems to derive from people who were already resident in Britain in 400 A.D., a substantial enough minority seems to have greater affinities to people who were living in the stretch of land between the Netherlands and Denmark.

The case for mass migration on the continent of Europe (with the exception of much of the Balkans) is more difficult to make in a cut & dried fashion because the basic outlines of Romanness were much more intact in the centuries after the fall than in Britain. Though France and Lombardy may have names which derive from German tribes, there is not much that is German about these regions today, and frankly, even at the height of the barbarian rule when conquest and migration were fresh, the non-Roman overlay was likely a thin elite layer. Outside of Britan and the Balkans, the languages of the Roman Empire and the Christian religion maintained their dominance even after the fall of the Roman political order, a transformation of social norms, and the collapse of the economy.

And yet this does not deny the possibility of migration of peoples into this order. In Empires and Barbarians: The Fall of Rome and the Birth of Europe the historian Peter Heather argues that we must not neglect the likelihood that to some extent the arrival of the Germans was one of “folk wanderings.” That the identity of the Franks, Goths, and Lombards, did not emerge ad hoc and de novo through the accrual of military men around a tiny nucleus of German warlords and their retainers. That women and children were also part of the movement into the Roman Empire. Heather, in fact, depicts the Gothic arrival as one of destitute refugees fleeing the famine and chaos outside of the Pax Romana, and their subsequent militarization and rebellion as one forced upon them by the exigencies of their situation.

A new preprint on bioRxiv, Understanding 6th-Century Barbarian Social Organization and Migration through Paleogenomics, clarify these arguments in the case of the Lombards, who conquered Italy in the 6th century. The abstract:

Despite centuries of research, much about the barbarian migrations that took place between the fourth and sixth centuries in Europe remains hotly debated. To better understand this key era that marks the dawn of modern European societies, we obtained ancient genomic DNA from 63 samples from two cemeteries (from Hungary and Northern Italy) that have been previously associated with the Longobards, a barbarian people that ruled large parts of Italy for over 200 years after invading from Pannonia in 568 CE. Our dense cemetery-based sampling revealed that each cemetery was primarily organized around one large pedigree, suggesting that biological relationships played an important role in these early Medieval societies. Moreover, we identified genetic structure in each cemetery involving at least two groups with different ancestry that were very distinct in terms of their funerary customs. Finally, our data was consistent with the proposed long-distance migration from Pannonia to Northern Italy.

The preprint has genetic and isotopic results from two graveyards associated with elite Lombards of the 6th century. The one in late antique Pannonia would be in modern Hungary. The one in modern Italy is near Turin. The late 6th century was a time of tumult in the Roman Empire, as both Italy and the Balkans were subject to massive turnovers of the ethnic and political orders. The movement into Italy from the northeast was a typical one, prefigured by the Goths and other Germans before the Lombards.

From what I know, as far as German barbarians went, the Lombards were rather “raw” and non-Roman (in contrast, some tribes, such as the Goths and Franks, had had relationships with the Roman Empire for generations before they decided to take it over). Though they were nominally Christianized, and elite Lombards persisted in practicing pagan rituals in Italy down to the 8th century, over 100 years after their conquest of the peninsula.

The authors used a lot of “best of breed” methods with their large data set, but the ADMIXTURE plot really illustrates the result fine enough. The blue is associated with Northwest European ancestry (British and white Utah samples), red with Italian ancestry (Tuscan), and green(ish) with Iberian (Spanish mostly). The very light blue is 1K Genomes Finnish. Panel B is the graveyard in modern Hungary, and panel C is the one from northern Italy.

There is a strong correlation in the graves with those being of Northern European ancestry, and having high status via grave goods. The individuals also exhibited some segregation in the graves. Northern European ancestry and Southern European ancestry individuals were clustered together. The Pannonian individuals, whether Northern or Southern European, don’t seem to resemble ancient or modern Hungarians. The isotope analysis indicates that many of the individuals were highly mobile.

Finally, the data was robust enough to do a pedigree analysis. It looks like a lot of these individuals are related. If you look at the plots you can see groups with the label “Kindred.”

There is so much detail in the results that I won’t recapitulate. Just read the preprint and make sure to check out the supplementary text. What I will say is this.

  1. The Lombard migration seems to have been a migration of people of Northwest European heritage into Southern Europe.
  2. The migration occurred during the lifetime of some individuals. These were highly mobile individuals.
  3. There were associated groups with the Lombards, who were genetically distinct, and likely of lower status. Their Southern European character is also distinct from the native population of Pannonia in the case of panel A.
  4. The Lombards themselves had Northern European ancestry which was somewhat heterogenous (probably different tribes and ethnicities). The shift away from Finnish ancestry probably indicates sampling more from western and opposed to central Europe.
  5. Admixture with the local populations and other post-Roman groups began early on.

The ethnocultural distinctiveness of the Lombards is clear from the textual evidence. The genetic data here confirm that in totality. But, The Geography of Recent Ancestry Across Europe, also highlighted a lot of deep population structure within modern Italy, and could not discern much impact of barbarian migration outside of the Balkans across their data set. Why?

It is rather clear that there were population declines across the West Roman Empire in the years after the Gothic Wars. If you read the textual evidence you imagine some sort of catastrophe going on. In human terms it was catastrophic. On the scale of economics, it was catastrophic. But in terms of population genetics, the long-term impact was not that extreme. The local population structure was not much altered because the Roman population base was so high that even a large decline did not induce bottleneck effects, and the German elite was also small enough it did not much perturb the underlying structure which had roots back to the period before the Roman Empire. Even in the first generations of Lombards in Italy, which is the Collego data set reflects, there was intermarriage between German people and others.

The demographic impact of the German migrations was huge on culture, politics, and economics. But it was not huge on population genetics.

May 11, 2017

When conquered pre-Greece took captive her rude Hellene conqueror

Filed under: Genetics,Genomics,Greece,History,Migration — Razib Khan @ 12:22 am


When I was a child in the 1980s I was captivated by Michael Wood’s documentary In Search of the Trojan War (he also wrote a book with the same name). I had read a fair amount of Greek mythology, prose translations of the Iliad, as well as ancient history. The contrast between the Classical Greeks, the strangeness of their mythology was always something that on the surface of my mind. The reality that Bronze Age Greeks were very different from Classical Greeks resolved this issue to some extent.

Though Classical Greeks were very different from us, to some extent Western civilization began with them, and they are very familiar to us. Rebecca Goldstein’s Plato at the Googleplex was predicated on the thesis that the ancient Greek philosopher had something to tell us, and that if he was alive today he would be a prominent public speaker.

I’m going to dodge the issue of Julian Jaynes’ bicameral mind, and just assert that people of the Bronze Age were fundamentally different from us. And that difference is preserved in aspects of Greek mythology. Though it is fashionable, and correct, to assert that Homer’s world was not that of Mycenaeans, but the barbarian period of the Greek Dark Age, it is not entirely true. Homer clearly preserved traditions where citadels such as Mycenae and Pylos were preeminent, and details such as the boar’s tusk helmets are also present in the Iliad.

But aesthetic details or geopolitics are not what struck me about Greek mythology, but events such as the sacrifice of Iphigenia. Like Abraham’s near sacrifice of his son, this plot element strikes moderns as cruel, barbaric, and unthinking. And though the Classical Greeks did not have our conception of human rights, they had turned against human sacrifice (and the Romans suppressed the practice when they conquered the Celts) on the whole, but it seems to have occurred in earlier periods.

The rupture between the world of the Classical Greeks and the strange edifices of Mycenaean Greece were such that scholars were shocked that the Linear B tablets of the Bronze Age were written in Greek when they were finally deciphered. In fact many of the names and deities on these tablets would be familiar to us today; the name Alexander and the goddess Athena are both attested to in Mycenaean tablets.

Preceding the Mycenaeans, who  emerge in the period between 1400-1600 BCE, are the Minoans, who seem to have developed organically in the Aegean in the 3rd millennium. This culture had relations with Egypt and the Near East, their own system of writing, and deeply influenced the motifs of the successor Mycenaean Greek civilization. The aesthetic similarities between Mycenaeans and Minoans is one reason that many were surprised that the former were Greek, because the Minoan language was likely not.

Mycenaean civilization seems to have been a highly militarized and stratified society. There is a reason that this is sometimes referred to as the “age of citadels.” Allusions to the Greeks, or Achaeans, in the diplomatic missives of the Egyptians and Hittites suggests that the lords of the Hellenes were reaver kings. In 1177 B.C. Eric Cline repeats the contention that a fair portion of the “sea peoples” who ravaged Egypt in the late Bronze Age were actually Greeks.

So when did these Greeks arrive to the shores of Hellas? In The Coming of the Greeks Robert Drews argued that the Greeks were part of a broader movement of mobile charioteers who toppled antique polities and turned them into their own. The Hittites and Mitanni were two examples of Indo-European ruling elites who took over a much more advanced civilizational superstructure and made it their own. While the Hittites and other Indo-Europeans, such as the Luwians and Armenians, slowly absorbed the non-Indo-European substrate of Anatolia, the Indo-Aryan Mitanni elite were linguistically absorbed by their non-Indo-European Hurrian subjects. Indo-Aryan elements persisted only their names, their gods, and tellingly, in a treatise on training horses for charioteers.

Drews’ thesis is that the Greek language percolated down from the warlords of the citadels and their retinues over the Bronze Age, with the relics who did not speak Greek persisting into the Classical period as the Pelasgians. Set against this is the thesis of Colin Renfrew that Greece was one of the first Indo-European languages, as Indo-European languages began in Anatolia.

The most recent genetic data suggest to me that both theses are likely to be wrong. The data are presented in two preprints The Population Genomics Of Archaeological Transition In West Iberia and The Genomic History Of Southeastern Europe. The two papers cover lots of different topics. But I want to focus on one aspect: gene flow from steppe populations into Southern Europe.

We know that in the centuries after 2900 BCE there was a massive eruption of individuals from the steppe fringe of Eastern Europe, and Northern Europe from Ireland to to Poland was genetically transformed. Though there was some assimilation of indigenous elements, it looks to be that the majority element in Northern Europe were descended from migrants.

For various reasons this was always less plausible for Southern Europe. The first reason is that Southern Europeans shared a lot of genetic similarities to Sardinians, who resembled Neolithic farmers. Admixture models generally suggested that in the peninsulas of Southern Europe the steppe-like ancestry was the minority component, not the majority, as was the case in Northern Europe.

These data confirm it. The Bronze Age in Portugal saw a shift toward steppe-inflected populations, but it was not a large shift. There seems to have been later gene flow too. But by and large the Iberian populations exhibit some continuity with late Neolithic populations.  This is not the case in Northern Europe.

In The Genomic History Of Southeastern Europe the authors note that steppe-like ancestry could be found sporadically during early periods, but that there was a notable increase in the Bronze Age, and later individuals in the Bronze Age had a higher fraction. Nevertheless, by and large it looks as if the steppe-like gene flow in the southerly Balkans (focusing on Bulgarian samples) was modest in comparison to the northern regions of Europe. Unfortunately I do not see any Greece Bronze Age samples, but it seems likely that steppe-like influence came into these groups after they arrived in Bulgaria, which is more northerly.

Down to the present day a non-Indo-European language, Basque, is spoken in Spain. Paleo-Sardinian survived down to the Classical period, and it too was not Indo-European. Similarly, non-Indo-European Pelasgian communities continued down to the period of city-states in Greece.

These long periods of coexistence point to the demographic equality (or even superiority) of the non-Indo-European populations. The dry climate of the Mediterranean peninsulas are not as suitable for cattle based agro-pastoralism. This may have limited the spread and dominance of Indo-Europeans. Additionally, the Mediterranean peninsulas were likely touched by Indo-European migrations relatively late. Much of the early zeal for expansion may have already dissipated by them. The high frequency of likely Indo-European R1b lineages among the Basques is curious, and may point to the spreading of male patronization networks, and their assimilation into non-Indo-European substrates where necessary. R1b is also found in Sardinia, and in high frequencies in much of Italy.

The interaction and synthesis between native and newcomer was likely intensive in the Mediterranean. For example, of the gods of the Greek pantheon only Zeus is indubitably of Indo-European origin. Some, such as Artemis, have clear Near Eastern antecedents. But other Greek gods may come down from the pre-Greek inhabitants of what became Greece.

Ultimately these copious interactions and transformations should not be a great surprise. The sunny lands of the Mediterranean attracted Northern European tribes during Classical antiquity. The Cimbri invasion of Italy, Galatians in Thrace and Anatolia, the folk wandering of Vandals and Goths into Iberia, are all instances of population movements southward. These likely moved the needle ever so slightly toward convergence between Northern and Southern Europe in terms of genetic content.

In relation to the more general spread of Indo-Europeans, I believe there are a few areas like Northern Europe, where replacement was preponderant (e.g., the Tarim basin). But I also believe there were many more which presented a Southern European model of synthesis and accommodation.

When conquered pre-Greece took captive her rude Hellene conqueror

Filed under: Genetics,Genomics,Greece,History,Migration — Razib Khan @ 12:22 am


When I was a child in the 1980s I was captivated by Michael Wood’s documentary In Search of the Trojan War (he also wrote a book with the same name). I had read a fair amount of Greek mythology, prose translations of the Iliad, as well as ancient history. The contrast between the Classical Greeks, the strangeness of their mythology was always something that on the surface of my mind. The reality that Bronze Age Greeks were very different from Classical Greeks resolved this issue to some extent.

Though Classical Greeks were very different from us, to some extent Western civilization began with them, and they are very familiar to us. Rebecca Goldstein’s Plato at the Googleplex was predicated on the thesis that the ancient Greek philosopher had something to tell us, and that if he was alive today he would be a prominent public speaker.

I’m going to dodge the issue of Julian Jaynes’ bicameral mind, and just assert that people of the Bronze Age were fundamentally different from us. And that difference is preserved in aspects of Greek mythology. Though it is fashionable, and correct, to assert that Homer’s world was not that of Mycenaeans, but the barbarian period of the Greek Dark Age, it is not entirely true. Homer clearly preserved traditions where citadels such as Mycenae and Pylos were preeminent, and details such as the boar’s tusk helmets are also present in the Iliad.

But aesthetic details or geopolitics are not what struck me about Greek mythology, but events such as the sacrifice of Iphigenia. Like Abraham’s near sacrifice of his son, this plot element strikes moderns as cruel, barbaric, and unthinking. And though the Classical Greeks did not have our conception of human rights, they had turned against human sacrifice (and the Romans suppressed the practice when they conquered the Celts) on the whole, but it seems to have occurred in earlier periods.

The rupture between the world of the Classical Greeks and the strange edifices of Mycenaean Greece were such that scholars were shocked that the Linear B tablets of the Bronze Age were written in Greek when they were finally deciphered. In fact many of the names and deities on these tablets would be familiar to us today; the name Alexander and the goddess Athena are both attested to in Mycenaean tablets.

Preceding the Mycenaeans, who  emerge in the period between 1400-1600 BCE, are the Minoans, who seem to have developed organically in the Aegean in the 3rd millennium. This culture had relations with Egypt and the Near East, their own system of writing, and deeply influenced the motifs of the successor Mycenaean Greek civilization. The aesthetic similarities between Mycenaeans and Minoans is one reason that many were surprised that the former were Greek, because the Minoan language was likely not.

Mycenaean civilization seems to have been a highly militarized and stratified society. There is a reason that this is sometimes referred to as the “age of citadels.” Allusions to the Greeks, or Achaeans, in the diplomatic missives of the Egyptians and Hittites suggests that the lords of the Hellenes were reaver kings. In 1177 B.C. Eric Cline repeats the contention that a fair portion of the “sea peoples” who ravaged Egypt in the late Bronze Age were actually Greeks.

So when did these Greeks arrive to the shores of Hellas? In The Coming of the Greeks Robert Drews argued that the Greeks were part of a broader movement of mobile charioteers who toppled antique polities and turned them into their own. The Hittites and Mitanni were two examples of Indo-European ruling elites who took over a much more advanced civilizational superstructure and made it their own. While the Hittites and other Indo-Europeans, such as the Luwians and Armenians, slowly absorbed the non-Indo-European substrate of Anatolia, the Indo-Aryan Mitanni elite were linguistically absorbed by their non-Indo-European Hurrian subjects. Indo-Aryan elements persisted only their names, their gods, and tellingly, in a treatise on training horses for charioteers.

Drews’ thesis is that the Greek language percolated down from the warlords of the citadels and their retinues over the Bronze Age, with the relics who did not speak Greek persisting into the Classical period as the Pelasgians. Set against this is the thesis of Colin Renfrew that Greece was one of the first Indo-European languages, as Indo-European languages began in Anatolia.

The most recent genetic data suggest to me that both theses are likely to be wrong. The data are presented in two preprints The Population Genomics Of Archaeological Transition In West Iberia and The Genomic History Of Southeastern Europe. The two papers cover lots of different topics. But I want to focus on one aspect: gene flow from steppe populations into Southern Europe.

We know that in the centuries after 2900 BCE there was a massive eruption of individuals from the steppe fringe of Eastern Europe, and Northern Europe from Ireland to to Poland was genetically transformed. Though there was some assimilation of indigenous elements, it looks to be that the majority element in Northern Europe were descended from migrants.

For various reasons this was always less plausible for Southern Europe. The first reason is that Southern Europeans shared a lot of genetic similarities to Sardinians, who resembled Neolithic farmers. Admixture models generally suggested that in the peninsulas of Southern Europe the steppe-like ancestry was the minority component, not the majority, as was the case in Northern Europe.

These data confirm it. The Bronze Age in Portugal saw a shift toward steppe-inflected populations, but it was not a large shift. There seems to have been later gene flow too. But by and large the Iberian populations exhibit some continuity with late Neolithic populations.  This is not the case in Northern Europe.

In The Genomic History Of Southeastern Europe the authors note that steppe-like ancestry could be found sporadically during early periods, but that there was a notable increase in the Bronze Age, and later individuals in the Bronze Age had a higher fraction. Nevertheless, by and large it looks as if the steppe-like gene flow in the southerly Balkans (focusing on Bulgarian samples) was modest in comparison to the northern regions of Europe. Unfortunately I do not see any Greece Bronze Age samples, but it seems likely that steppe-like influence came into these groups after they arrived in Bulgaria, which is more northerly.

Down to the present day a non-Indo-European language, Basque, is spoken in Spain. Paleo-Sardinian survived down to the Classical period, and it too was not Indo-European. Similarly, non-Indo-European Pelasgian communities continued down to the period of city-states in Greece.

These long periods of coexistence point to the demographic equality (or even superiority) of the non-Indo-European populations. The dry climate of the Mediterranean peninsulas are not as suitable for cattle based agro-pastoralism. This may have limited the spread and dominance of Indo-Europeans. Additionally, the Mediterranean peninsulas were likely touched by Indo-European migrations relatively late. Much of the early zeal for expansion may have already dissipated by them. The high frequency of likely Indo-European R1b lineages among the Basques is curious, and may point to the spreading of male patronization networks, and their assimilation into non-Indo-European substrates where necessary. R1b is also found in Sardinia, and in high frequencies in much of Italy.

The interaction and synthesis between native and newcomer was likely intensive in the Mediterranean. For example, of the gods of the Greek pantheon only Zeus is indubitably of Indo-European origin. Some, such as Artemis, have clear Near Eastern antecedents. But other Greek gods may come down from the pre-Greek inhabitants of what became Greece.

Ultimately these copious interactions and transformations should not be a great surprise. The sunny lands of the Mediterranean attracted Northern European tribes during Classical antiquity. The Cimbri invasion of Italy, Galatians in Thrace and Anatolia, the folk wandering of Vandals and Goths into Iberia, are all instances of population movements southward. These likely moved the needle ever so slightly toward convergence between Northern and Southern Europe in terms of genetic content.

In relation to the more general spread of Indo-Europeans, I believe there are a few areas like Northern Europe, where replacement was preponderant (e.g., the Tarim basin). But I also believe there were many more which presented a Southern European model of synthesis and accommodation.

April 8, 2017

Why only one migrant per generation keeps divergence at bay

The best thing about population genetics is that because it’s a way of thinking and modeling the world it can be quite versatile. If Thinking Like An Economist is a way to analyze the world rationally, thinking like a population geneticist allows you to have the big picture on the past, present, and future, of life.

I have some personal knowledge of this as a transformative experience. My own background was in biochemistry before I became interested in population genetics as an outgrowth of my lifelong fascination with evolutionary biology. It’s not exactly useless knowing all the steps of the Krebs cycle, but it lacks in generality. In his autobiography I recall Isaac Asimov stating that one of the main benefits of his background as a biochemist was that he could rattle off the names on medicine bottles with fluency. Unless you are an active researcher in biochemistry your specialized research is quite abstruse. Population genetics tends to be more applicable to general phenomena.

In a post below I made a comment about how one migrant per generation or so is sufficient to prevent divergence between two populations. This is an old heuristic which goes back to Sewall Wright, and is encapsulated in the formalism to the left. Basically the divergence, as measured by Fst, is proportional to the inverse of 4 time the proportion of migrants times the total population + 1. The mN is equivalent to the number of migrants per generation (proportion times the total population). As the mN become very large, the Fst converges to zero.

The intuition is pretty simple. Image you have two populations which separate at a specific time. For example, sea level rise, so now you have a mainland and island population. Since before sea level rise the two populations were one random mating population their initial allele frequencies are the same at t = 0. But once they are separated random drift should begin to subject them to divergence, so that more and more of their genes exhibit differences in allele frequencies (ergo, Fst, the between population proportion of genetic variation, increases from 0).

Now add to this the parameter of migration. Why is one migrant per generation sufficient to keep divergence low? The two extreme scenarios are like so:

  1. Large populations change allele frequency very slowly due to drift, so only a small proportion of migration is needed to prevent them from diverging
  2. Small populations change allele frequency very fast due to drift, so a larger proportion of migration is needed to prevent them from drifting

Within a large population one migrant is a small proportion, but drift is occurring very slowly. Within a small population drift is occurring fast, but one migrant is a relatively large proportion of a small population.

Obviously this is a stylized fact with many details which need elaborating. Some conservation geneticists believe that the focus on one migrant is wrongheaded, and the number should be set closer to 10 migrants.

But it still gets at a major intuition: gene flow is extremely powerful and effective at reducing differences between groups. This is why most geneticists are skeptical of sympatric speciation. Though the focus above is on drift, the same intuition applies to selective divergence. Gene flow between populations work at cross-purposes with selection which drives two groups toward different equilibrium frequencies.

This is why it was surprising when results showed that Mesolithic hunter-gatherers and farmers in Europe were extremely genetically distinct in close proximity for on the order of 1,000 years. That being said, strong genetic differentiation persists between Pygmy peoples and their agriculturalist neighbors, despite a long history of living nearby each other (Pygmies do not have their own indigenous languages, but speak the tongue of their farmer neighbors). In the context of animals physical separation is often necessary for divergence, but for humans cultural differences can enforce surprisingly strong taboos. Culture is as strong a phenomenon as mountains or rivers….

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