Ancient DNA Provides A New Understanding of Haplogroup I2a1a M26

Below is a map of confirmed instances of I-M26 found in prehistoric remains.  Lots of others could have potentially been added -- ancestral clades, closely related sister clades, and ones where the coverages is insufficient to determine whether it is indeed M26 (or L672 L160 etc.)  But we decided to err on the side of being conservative.

Going chronologically, we have M26 in what is now Sweden, at the Motala site, at 7730 BP (Before Present).  These were Hunter/Gatherers.

Next, we have it at the La Spina site, in modern Spain, at 5765 BP.  This was a Megalithic site, during the Neolithic.  Farming was already in Spain at that time.

Next we find it at Treilles, modern France, at 5015 BP.  This was a Megalithic/Neolithic site of farmers, near the coast.

Next we find it in the Remedello culture, of Northern Italy, at 4758 BP.  This was a Chalcolithic (Copper Age) site.  Per Robert S.P. Beekes, "bears all the marks of an Indo-European invasion: a new style of ceramics, a new burial rite, changes in the social structure, the introduction of a warrior aristocracy, the intro- duction of metallurgy, the horse and the chariot. But it is still not possible to assign language groups to particular culture."

Finally, for now, researchers have found it in the Megalithic culture of central France, near the Dolmen of Villaneuve-Sur-Yonne, 4753 BP, and again, of course, farming was present.  

In modern times, we find M26 at 37% in Sardinia, certainly an outlier due to substantial founder effect.

But it is also found at up to 10% in Samnite country in Molise, Italy (and in significant numbers in Cosenza province of northern Calabria).  It is found at 5-9% in Spain, including Basque country.  At up to 7% on Sicily.  And 3% in many areas of England and Ireland, especially places like the Channel Islands.  It is still found at 1% in Southern Sweden.

So, given all that we know, what is a sensible theory for M26's distribution and spread?  

We've heard them all, and each has merit: that is represents Megalithic Mariners, who went around old Europe converting local populations and building huge monuments like Stonehenge and the Nuraghe.   That it represents the spread of Cardial Ware culture, along the western Mediterranean seaboard.  That it represents the spread of farming, either as hunter/gatherers who adopted farming quickly, or as a rare clade of Haplogroup I that was predominantly farmers since the dawn of agriculture.  Finally, some posit that it represents a caste (or not) of people embedded in other haplogroups in motion, most often listed as G2a or R1b.

All have merit, none are perfect.  Let's go through the logical conclusions and form a model.

The presence of M26 amongst the hunter/gatherers of Motala, and its widespread ancient distribution by the dawn of the Neolithic tell us that it is a most ancient indigenous European clade.  

We believe that it is safe to say that the first modern humans in Europe, Aurignacians, aka Cro-Magnons, bore haplogroup C.  (Although during the Paleolithic, that far back, it is really anyone's guess whether Hg C came in during one the next phases).

M26, along with its brother clades within Haplogroup I2, most likely formed a part of the second wave of European hunter/gatherers, and was presumably present among the Epi-Gravettians and the Magdalenians.

How then does one explain its wide distribution and adaptive nature throughout the continent during the Neolithic? 

Perhaps the answer is simple.  These were people who have been in Europe for a long time, and are adaptable, and are survivors.

Europe was settled in waves, and not just the three big ones (Hunter/Gatherer, Farmers, Steppe Horsemen), which is an oversimplification.  

Imagine Europe as a pipe.  It has three entry points: the steppes of Russia from people heading due west, the Balkans/river corridors from people heading northwest, and the Mediterranean from people heading north.

Imagine a party.  The "Emtwentysix" family was among the first to arrive.  When they got to the house, they represented maybe 15% of the guests.  But as more guests arrived, some through the front door, some through the back door, and some through the side door, the Emtwentysix family became a smaller percentage of the total guests, now just 0.5-3%.  

When the family arrived, they were playing foosball together.  But at some point, some members of the family joined a group of other guests who were dancing, and some joined groups of other guests who were playing video games.

You get the analogy.  

None of the above theories may be 100% accurate, and yet all theories may be accurate in their own way. 

For example, it is doubtful that I-M26 first showed up in Europe with the Cardium Pottery culture, but it could have been among the groups (as it was clearly in Italy for a long time) that encountered the Cardial peoples, and then became a component of said peoples, heading west along the western Mediterranean seaboard.  

Similarly, in Northern Italy, M26 peoples could have been among the first who were Indo-Europeanized, in the Remedello culture, and then part of the secondary expansion into Italy, which formed the Oscan-speaking Sabellic tribes (Samnites, Brutti, and Sicels).  

In other places, clearly M26 was Megalithic acculturated, and its odd distribution in places where Megaliths appear is intriguing for sure.  

And lastly, some of its distribution could reflect later movements by obsidian traders or something similar.

Prehistory is a series of periods of demographic expansion followed by demographic crisis.  Good hunting, good weather, good crops, absence of disease, and other factors make people have more babies.  Then luck turns, and some lines die out, while other lines come to the party.  Since the Bronze Age in Europe, it has mostly been a demographic march to more population as time goes by.  

We would like to see a study comparing the M26 in Sweden, Ireland, Spain, the Italian mainland, and Sardinia, to see who is ancestral to whom, or how and when the different groups separated.

But the bottom line appears to be that M26 has been in Europe a long time, and like all lineages that were there a long time, its distribution will have changed a bit and its absolute numbers will have gone down, but these Most Adaptable Hunter Gatherers (MAHGs) continue to intrigue.




 

Allentoft Study Shows I2a1a in Bronze Age Remedello in Northern Italy

If you've been following Ancient DNA lately, you know that Allentoft et al came out with results of eastern Bronze Age populations.  The data confirm an expansion out of the steppes and likely give credence to a two or three wave Indo European expansion.

Paul Heggarty gives an excellent analysis here, perhaps the best on the web.

This post will focus on the study's implications for I-M26.  It is possible that M26 was linked perhaps to the spread of Cardium Pottery, which spread from Italy rapidly: 2,000 km from the gulf of Genoa to the estuary of the Mondego in probably no more than 100–200 years. This suggests a seafaring expansion by planting colonies along the coast.

Whether M26 spread with Cardial Ware, or existed before it and those people became acculturated by it, with the findings from Treilles and Remedello, we have confirmation that M26 was the lineage of some of the very first waves of inhabitants of Italy and the western Mediterranean, and that it persisted through very different times, since the Remedello culture was very different from its predecessors and very similar to steppe cultures.

How Little We Know About Ancient DNA

I've frequented several of the Ancient DNA discussion boards lately, and have been flummoxed by the self-important, self-promoted, self-described "experts," who proclaim to know precise migration patterns of Ancient Europeans.

These same "experts" even go so far as to claim to be able to tie specific haplogroups to languages, tribes, and epochs.  They will make broad statements, like, "all of Europe was populated by [this haplogroup or that], which represented the [Cro-Magnons or whatever], until they were replaced, en masse, by the [new Haplogroup.]"

(Often the dominant invader haplogroup in their theories tends to be the one of the posting "expert," but that's just coincidence, I'm sure.)

Contrasting these experts are some bona fide theoreticians, who point out that we have less than 100 samples of Caucasian Ancient DNA, and that a simple cultural fact, for example, if one tribe cremated their dead and another tribe buried their dead, could contribute to the number of ancient skeletons that make it to the present day.

So, what I decided to do was to plot the confirmed ancient NR Y Chromosome haplogroup samples on a map, to show whatever it shows.

What I discovered was a complete lack of any real patterns.  In other words, it's too early to tell.  We need way more aDNA.

I used the excellent data from Ancestral Journeys.  All maps are labeled.  All times and locations are approximate.  All maps are copyrighted, but feel free to share, as long as you link to this page or attribute to me.  (The final map is not mine, but purports to represent modern majorities).

I think from these maps it is clear that several of the widely accepted theories are bunk.  For example, looking at these maps, it is clear that Haplogroup G2 is a candidate too for one of the original populations of Europe.  It was ubiquitous.  The wiseguys all postulate that it originated in the western Caucasus, near where it is currently dominant, and moved west with the migratory herders or agriculturalists.  However, it is just as likely from looking at these maps that it once simply was everywhere, in a band in central Europe, along the major rivers, stretching from northern Spain to the Caucasus, and that its current location is one where it RECEDED to, not originated from.

What other theories can be questioned by these maps?

Corrected European Regional Percentages (Yet Again) for I-M26 12a1a

Some have pointed out that the early studies showing high M26 in Castile suffered from small sample sizes.  Further, more recent posts and agglomerations of studies show different numbers than what is quoted below.  While Eupedia is usually full of pseudoscientific malarky, this post purports to have collected studies.  And this website claims to have done the same, specific to Spain.

So, taken together, we get the following on our M26 I2a1a leaderboard:

(1) Sardinia (Island Isolated) ~37%
(2) Samnium (Mountainous Central Italy) ~10%
(3) Navarre (Mountainous Northern Spain) ~9%
(4) Sicily (Another Island) ~7%
(5) Basque Country (More Mountains) ~5%
(6) Aragon, Spain ~5%
(7) England, Ireland, and ALL W. European Islands whether Mediterranean, Atlantic, or Channel  ~3%


We also have some allegations (no studies cited) that Andalusia (Spain) is in double digits.


From what I can tell, this is the picture that has emerged.  Please cite links to studies if you have any.

Corrected Study Confirms Even More I-M26 12a1a In Archaic Zones of Italy

The great paper by Brisghelli, et. al., referred to and discussed extensively below, has been updated and corrected.  The updates are stunning.  Now, the descendants of the once-mighty Samnites, the rugged mountain-dwellers of the Southern Italian interior, have shown to be 10% I-M26.

The corrected table can be found by clicking this link.

This pushes Samnites into third place in the M26 frequency chart.  It also confirms what I've posted previously about M26 being found in large numbers among Northern Calabrian mountaineers, who have long been noted to be offshoots of the Samnites.  (The Ancient sources said that the Brutti broke off from the Lucanians, and the Lucanians broke off from the Samnites.  Each of these hardy tribes battle Rome for supremacy in pre-Empire Italy, and they almost won.)

But I digress.  Here are the updated charts of frequency for M-26:


Sardinia ~37%
Castile ~19%
Samnium ~10%
Sicily ~7%
Basque Country ~5%
England, Ireland, and ALL W. European Islands whether Mediterranean, Atlantic, or Channel  ~3%
General Italy/France ~2%


This might clear things up a bit for the spread of M26.

M26's ancestral clade, P37.2, is concentrated very strongly in the Balkans, and that all data seems to indicate that M26 followed a route that went westward through Italy.  In other words, if there had been no stop in Italy, there'd be no stop in Sardinia.  No Italy, no southern France.  Thus, unless there was a back migration of M26 eastward, the M26 there is likely to be ancestral to the M26 that is more westerly. 

An eastward migration is highly unlikely, be cause the Italian regions where it is found (South and Central Appenines) are extremely isolated, and far from the typical coastal routes.  So it is unlikely that a few Sardinian slaves, for example, or English pirates or whatever, spread it.  Samnium and Cosenza are way up in the mountains.

Also, there is a U5 mtDNA clade in Sardinia connected to M26 that bears evidence of an Italian refugium that contributed (in a small way) to the post-glacial expansion of mankind into Europe. 

In other words, synthesizing the various sources, one gets: I2 likely evolved in Anatolia a very long time ago.  It spread westward for sure.  A major stopping point was the Balkans, where regular old P37.2 evolved.  Another stop was Italy, likely, perhaps Piedmont, which is on the French border.  M26 likely evolved there, perhaps during a period of glacial expansion.

Alternatively, the timing of this westward spread is later, and mirrors the Cardium culture.  Either way, South Central Italy (Samnium) is directly west of the Balkans.  And it is isolated.  A strong candidate for a similar Founder effect, like what happened in Sardinia. 

From Italy, it spread to Sardinia, and points further west.  M26 then had a second expansion during the Bronze or Copper Age, which was seaborne.  Others have speculated this mirrors the Megalith culture, and I see no reason why this is not a possibility.  The second spread was clearly seaborne.

I think the M26 found way up in the remote mountains of Italy is a great candidate to be ancestral to the others.  We need some more research on the SNPs to solve this once and for all.  As recent R1b research has shown, the STRs are not enough.

Scholars Finally Apply Some Logic (Not Mythology) To Etruscan Origins

If you read the old Roman and Greek historians, you will notice that they mix their religions and mythology into their writing.  For example, a casual perusal of Suetonius will reveal detailed historical facts and stuff about how talking cows portended the eventual rise of this Emperor or that.  Even the most serious historians will pass along foundation myths, for example, how Hercules visited this German tribe or that, or how some random tribe of Spaniards or Gauls claimed descent from Troy.

No one takes these other facts seriously, except with respect to the Etruscans.

It's been well-documented that the Etruscans (and the Romans) both embellished ties to the older civilizations of the east, only upon an uptick in contacts with the East. 

The Romans invented ties with Troy to beef up their bona fides when faced with older, Hellenistic cultures.  The Etruscans did this too.

When the Etruscans were battling the Greeks for cultural supremacy, they constantly faced snobbery because the Greeks were "older" than them.  So the Etruscans, who also wanted increased trade with Asia Minor, invented ties with the Lydians there.  One or two ancient historians reported this, and enshrined the myth.

If knowledge of Ancient History ain't your thing, just consider how many modern rappers of African-American heritage (to establish their bonafides) call themselves "Gotti" or some variation of the Italian Godfather theme. 

The problem is that certain modern armchair historians, due to lack of perspective, or some animus toward how much of Western Civ originated in Italy, or due to some notion that Northern Italians racially superior, welcome and embrace the exotic Etruscans hypothesis.  They have done so for a while, despite absolutely ZERO evidence of any invasion or cultural shift in Etruscan areas in pre-Etruscan times.  In fact, there is nothing but continuity between Villanovans and their Etruscan successors in the region!

Anyway, now we have genetic studies that finally shut the door on the dummies. 

The link can be found here. 

The study basically found that there are no more ties with Etruscans to the Near East than there are for other Mediterranean cultures, and that any such links are due to the very first farmers migrating to populate Europe at the dawn of post-Ice Age history.

This has been been confirmed by other evidence too, summarized here.

Can we finally put this "Exotic Etruscans" myth to bed, alongside the other ancient myths, like that Aeneas carried the founders of Rome to Italy from Troy on his back, and that eagles pooping caused Vespasian to become Emperor?  Those also appear in ancient histories, after all.

EXCITING NEWS: New Study Finds Significant M26 Among South Central Italy Mountain Dwellers

Confirms theory that M26 likely represents aboriginal Italians.

In 2009, Maria Pala et al first put out the theory that YCC Hg I-M26 represented the aboriginal Italians, who had stayed in a small Italian refugium during the LGM.  The paper was called "Mitochondrial Haplogroup U5b3: A Distant Echo of the Epipaleolithic in Italy and the Legacy of the Early Sardinians" and can be found here.  It was kind of an added bonus theory in the paper, which focused on mtDNA but put forth the idea en passant.

Now, a landmark study on Italian Y chromosomes shows significant I-M26 in the archaic Samnium region of Italy.  This is and was always a rugged, remote, mountainous part of Italy known even in Roman times as being populated by some of the earliest inhabitants of Italy.  This, coupled with what American genetic genealogists already knew (which is the presence of M26 in many males from remote parts of mountainous northern Calabria), would seem to confirm the theory that M26 was present in an Italian refugium during the epipaleolithic.  And that those people went on to populate Sardinia and other coastal regions of southwest Europe.

Is Y-Chromosome Haplogroup M26 Norman? The Answer Can Be Found By Resorting to Logic

M26 is a clade of Haplogroup I on the Y Chromosome.  (It is sometimes called I2a1, but I disfavor such nomenclature, because I have lived through M26 being renamed often.  It is subject to subjective and regular change, which makes research hard.  M26 used to be known as Eu-8 and I1b2.) 

M26 is found in large numbers in Sardinia.  Between 37-40% of the males there bear it.  It is also found regularly in the Iberian peninsula: Basque Country, Castile, Aragon, Portugal, etc.  Lastly, all of the islands in Western Europe show M26 as well: Ireland, England, the Balearic Islands, Pantellaria, etc.  Some have stated that M26 must have spread, after its initial emergence, via the ocean, since it is found almost exclusively along the coasts of western Europe.

There are tens of theories on what the spread of M26 correlates to, both prehistoric and historical.  Some posit it tracks the spread of Cardium Pottery or Atlantic Bronze Age culture or Megalith builders.  Many now believe that M26 tracks Sardinian obsidian traders, because (a) it has been shown that Sardinian obsidian traders sailed the oceans plying their wares; (b) during that period was the only period where Sardinia was an exporter of humans; (c) and of course the large concentration of M26 in Sardinia.  All of these theories relate to prehistoric happenings, of course. 

Some believe M26 tracks historical migrations.  Possible candidates include the Roman slave trade, the Spanish empire, or the Normans. 

This exercise attempts to go through each theory as it relates to the discovery of several I-M26 in the Calabria region of South Italy.  M26’s presence in Calabria could likely be the key to resolving the issue once and for all, yet this seems to have escaped modern scholarship.

Below is the list of just a few of the publicly available Calabrian clans that bear M26, tested by a major lab:

Caracciolo – exact origin unknown

Faga – Cortale, Catanzaro

Funicella – San Vincenzo La Costa, Cosenza

Gatto – Lago, Cosenza

Lamantia – Sicilian, but originally from Amantea, Cosenza

Rondinella – Gioiosa Ionica, Reggio Calabria

The presence of M26 in Calabria is rather unique, since it is the easternmost that it is found in real quantity.  It also is not an obvious place for it.   

Below, each theory for M26 in Calabria is stated, and how this fits in the puzzle, is examined.  Then, I offer an opinion on each theory.

(1) There is no significance to Calabrian M26:  Lots of Calabrians have tested M26 because DNA testing is largely an American and UK phenomenon, and Calabrians simply represent a large portion of Italian-Americans.  As a threshold, this theory cannot be ignored.  As anyone who participates in genetic genealogy knows, there are far more samples of R1b because home DNA testing has lots of western European participants.  Under this theory, M26 would have a steady distribution across most of Italy, but would show up in so many American genetic genealogy sites simply because a large percentage of those getting tested were Italian-Americans from Calabria.  Evaluation: unlikely.  There is a large corpus of works testing Italians in Italy, and M26 has shown up in several regions besides Calabria.  But if M26 were indeed evenly spread across Italy, we would expect to see large numbers of test kits coming back M26 from other large groups of Italian-American source populations, i.e., Neapolitans.  This has not been the case.  There do appear to be higher percentages of M26 in Calabria.

(2) Calabrian M26 signifies the first humans in Italy:  Italy was a small refugium during the LGM and the males who existed there were M26.  M26 spread along the coast of Italy and found its way to Sardinia, where it had a founder effect.  Calabria, as the toe of Italy and with its inpenetrable mountains, is where the remnants of a relict M26 population were driven by subsequent invaders.  M26 thus represents aboriginal Italians from way back.  This theory was set out in “Mitochondrial haplogroup U5b3: a distant echo of the epipaleolithic in Italy and the legacy of the early Sardinians.”  (Available for free in its entirety on Google).  It is buttressed by the earliest Homo Sapiens and significant Gravettian finds being in the Calabria area.  (Grotto della Romito, etc.)  It also would make sense that aboriginal Italians would be slowly driven into the wild mountains at the very bottom of the peninsula, by subsequent invasions over time.  Evaluation: very possible.  There is an easy way to test this.  Either the Calabrian M26 represents the foundational (upstream) branch, or it doesn’t.  Either it shows itself to be ancestral to Sardinian M26 (and all others), or it does not.  Also, Calabrian M26 samples should be very divergent from those in the rest of Western Europe.

(3) M26 was spread WEST by the cardial ware culture.  Makers of Cardium pottery have been shown to have taken a western route along the Mediterranean, and cardial ware has been posited to correlate with M26.  The M26 is Calabria represents those who stayed behind in Italy on the way to Sardinia.  This Evaluation: possible.   Again, there is an easy way to test this.  Either the Calabrian M26 represents the foundational (upstream) branch, or it doesn’t.  Either it shows itself to be as old as Sardinian M26, or it does not.

(4) M26 was spread EAST:  M26 was spread easterly into Italy, from more western Atlantic Bronze Age cultures, or from Sardinian obsidian traders, or from Western European Megalith builders.  Again, if you Google search or Wikipedia these concepts, you can readily see the various theories.  Sardinian obsidian, the black gold (or more accurately, the advanced military hardware) of ages past, was spread by Sardinian obsidian traders all over western Europe, for one example.  Evaluation: possible.   As above, there is an easy way to test this.  The Calabrian M26 should show its closest MRCA to be that of Sardinian M26, and this should be between 3000 and 4000 years ago.

Possible Historical Sources:

(5) Roman era causes:  M26 in Calabria represents Sardinian slaves brought into Calabria during Roman times.  Rome fought a series of wars in Sardinia around 200 B.C.  They brought back vast quantities of slaves.  For a while, Romans had a saying, “as cheap as a Sardinian slave.”  Rome also conquered Calabria around this time, and turned vast tracts into public lands.  The public lands were cultivated by slaves, and their descendants bear M26.  If ~40% of Sardinian slaves bore M26, it follows that a few propagated, and entered the general population.  Evaluation: possible but not likely.  There were many other agricultural regions in Italy besides Calabria.  If Sardinian slaves were so prevalent around Italy in 200 BC, there should be an even spread of M26 in Italy.  There simply is not.  Yes, there are other mechanisms (Iberian mercenaries in Hannibal’s army, Spartacus revolts) that could explain a Roman era M26 propagation.  But again, these are easily testable.  If Calabrian M26 represents Sardinian slaves from 200 BC (or Iberian mercenaries from the same time), those who tested positive should cluster very closely with the source populations.  I don’t believe they do.

(6) Normans:  M26 in Calabria represents the blood of Viking overlords.  The Normans did by coincidence conquer a number of lands where there is M26: the UK, Calabria, etc.  M26 in Calabria could represent the descendants of these individuals.  Evaluation: unlikely.  The Normans also didn’t conquer many lands where there is M26.  They had no presence in Basque country, many of the Spanish states, the interior of Ireland, Pantellaria in Italy, etc.  Plus the Calabrian M26 would need to cluster with Northern French M26, which it does not.  Furthermore, the Norman invasion of Italy was only about 1000 years ago, not only during historical times, but at the cusp of surnames.  I believe most of those testing positive for M26 from Calabria do not bear Norman surnames.  (Examples of Norman surnames are Ruggiero, Gandolfini, etc.)  Those testing positive for M26 from Calabria have Latin surnames.

(7) Spaniards:  The Spanish ruled Calabria so the M26 there likely just comes from Spanish settlers.  There is much M26 is certain provinces of Spain and therefore, if 20% of colonists, mercenaries, or adventurers bore the gene, it would make sense that a percentage of Calabrians still bear the gene today.  Evaluation: unlikely.  Again, easily testable:  The Spanish presence in Calabria was so recent that Calabrians testing positive for M26 should be virtually identical to Spanish sources.  It’s not.  Moreover, some of the Calabrian families testing positive for M26 have their trees traced back to the 1500s, with only Italian names. 

Taking the big picture here, you can decide.  I personally think Theory (2) or (3) is most likely, that M26 in Calabria represents a very ancient people.  Even in Roman times, Calabria was called the First Italy.  The tribes that existed there that blended with the Romans were called the Itali (from which the entire peninsula was named).  The mountains in Calabria, with dense pine forests, high peaks covered with snow, and non-navigable passes are an ideal place for the original Italian population to be found. 

Comparing the STR data from these  Calabrian families, it shows that they are most closely related to other Calabrians, and estimates of TMRCA among the Calabrians is as recent as 500 years ago.  The next closest from what I could see from a very quick look were samples from the UK, England and Ireland, with a TMCRA back in prehistory.  This would seem to eliminate theories 5-7 above.