jueves, 15 de septiembre de 2011

Y haplogroup R1b1a2a1a1b (S116+)

44 Certified STR Markers

DYS19 / 394 = 13
DYS385a = 11
DYS385b = 14
DYS388 = 12
DYS389i = 13
DYS389ii = 29
DYS390 = 24
DYS391 = 11
DYS392 = 14
DYS393 / 395 = 13
DYS426 = 12
DYS437 / 457 = 14
DYS438 = 12
DYS439 / GATA A4 = 12
DYS442 = 12
DYS445 = 12
DYS446 = 13
DYS447 = 25
DYS448 = 18
DYS449 = 30
DYS453 = 11
DYS454 = 11
DYS455 = 11
DYS456 = 16
DYS458 = 18
DYS459a = 9
DYS459b = 10
DYS460 / GATA A7 1 = 11
DYS461 / GATA A7 2 = 12
DYS462 = 11
DYS464a = 14
DYS468 = 29
DYS484 = 13
DYS522 = 10
DYS527a = 20
DYS527b = 21
DYS531 = 11
DYS557 = 16
DYS588 = 18
GATA A10 = 13
GATA C4 = 24
GATA H4 = 11
YCA IIa = 19
YCA IIb = 23YS438 12
DYS439 12
DYS447 25
DYS448 18
DYS460 11
GATA H4 11
YCAIIa 19
YCAIIb 23
===============

Timeline of Y-DNA Haplogroup R1b -- BT SRY10831.1 (+) CF M168 (+) C+F -- F M89 (+) IJK -- K M9 (+) P M45 (+) R M207 (+) R1 M173 (+) R1b M343 (+)

Haplogroup R1b
Time: approx 10,000 years ago
Place: Western Europe
Facts: The founder of the R1b lineage lived over 35,000 years ago prior to the end of the last Ice Age in southern Europe and Iberia.

Members of Haplogroup R1b are believed to be descendants of Cro-Magnon people, the first modern humans to enter Europe.

Cro-Magnons lived from about 35,000 to 10,000 years ago in the Upper Paleolithis period of the Pleistocene. Cro-Magnons were anatomically modern, differing from their modern day descendants in Europe by their slightly more robust physiology.

Cro-Magnons buried their dead intentionally, and likely had a knowledge of ritual, by burying their dead with necklaces and tools. Surviving Cro-Magnon artifacts include huts, cave paintings, carvings and antler-tipped spears.

It is believed that the Cro-Magnon came into contact with the Neanderthals, who inhabited Europe 230,00 to 29,000 years ago and is thought to have caused the extinction of the Neanderthals. There is also recent fossil evidence that the Cro-Magnon people interbred with the Neanderthals.

When the ice sheets retracted at the end of the ice age, descendents of the R1b lineage migrated throughout western Europe.

Today, Haplogroup R1b is found predominantly in western Europe, including England, Ireland, and parts of Spain and Portugal. It is especially concentrated in the west of Ireland where it can approach 100% of the population.

This haplogroup contains the well known Atlantic Modal STR Haplotype (AMH). AMH is the most frequently occurring haplotype amongst human males with an Atlantic European ancestry. It is also the haplotype of Niall of the Nine Hostages, an Irish King in the Dark Ages who is the common ancestor of many people of Irish patrilineal descent.

My Paternal Genome -- From Genebase
(sample contributed by my brother Ismael H Camacho)
The Camacho genome ancestry map (Click here for map of all migrations)

Type: Y-Chromosome
DNA sample donated by Ismael H Camacho

Your Y-chromosome results identify you as a member of haplogroup R1b.

SNP Location SNP Identity Result Mutation
L62 A > G A Negative
M11 A > G A Negative
M122 T > C T Negative
M130 C > T C Negative
M168 C > T T Positive
M170 A > C A Negative
M173 A > C C Positive
M174 T > C T Negative
M175 TTCTC del No TTCTC del Negative
M201 G > T G Negative
M207 A > G G Positive
M214 T > C T Negative
M230 T > A T Negative
M231 G > A G Negative
M242 C > T C Negative
M304 A > C A Negative
M343 C > A A Positive
M45 G > A A Positive
M60 T ins No T ins Negative
M69 T > C T Negative
M70 A > C A Negative
M89 C > T T Positive
M9 C > G G Positive
M91 T del No T del Negative
M96 G > C G Negative
P256 G > A G Negative
SRY10831.1 A > G G Positive
SRY10831.2 G > A G Negative
=======================================

Based on Y-DNA R Subclade Test, Ismael Hernando's Y-DNA subclade has been confirmed as R1b1a2a1a1b (S116+).
SNP Location SNP Identity Result Mutation
M124 C > T C Negative
M126 AATA del No AATA del Negative
M153 T > A T Negative
M157 A > C A Negative
M160 A > C A Negative
M167 C > T C Negative
M18 AA ins No AA ins Negative
M198 C > T C Negative
M222 G > A G Negative
M269 T > C C Positive
M335 T > A T Negative
M37 C > T C Negative
M56 A > T A Negative
M65 A > T A Negative
M73 GT del No GT del Negative
M87 T > C T Negative
P107 G > A G Negative
P25 C > A A Positive
P297 G > C C Positive
P66 G > A G Negative
P98 C > T C Negative
PK5 C > T C Negative
S116 C > A A Positive
S127 T > C C Positive
S21 C > T C Negative
S26 A > C A Negative
S28 G > A G Negative
S29 G > A G Negative

=====================================

Interpretation of the Results:

The above results are from the laboratory analysis of the Camacho Y-chromosome. The DNA was analyzed for Short Tandem Repeats (STRs), which are repeating segments of the genome that have a high mutation rate.

The location on the Y chromosome of each of these markers is depicted in the image (to the right), with the number of repeats for each of the STRs presented to the right of the marker (above). For example, DYS19 is a repeat of TAGA, so if the DNA repeated that sequence 14 times at that location, it would appear: DYS19 14.

Studying the combination of these STR lengths in the Y Chromosome allows researchers to place the subject in a haplogroup, which reveals the complex migratory journeys of the ancestors.

The Camacho Y-chromosome results identify the Camachos as a member of haplogroup R1b. The genetic markers that define this ancestral history reach back roughly 60,000 years to the first common marker of all non-African men, M168, and follow the lineage to present day, ending with M343, the defining marker of haplogroup R1b.







Looking at the map highlighting the Camacho ancestors' route, the members of haplogroup R1b carry the following Y-chromosome markers:
M168 > M89 > M9 > M45 > M207 > M173 > M343

Today, roughly 70 percent of the men in southern England belong to Haplogroup R1b. In parts of Spain and Ireland, that number exceeds 90 percent.

What's a haplogroup, and why do geneticists concentrate on the Y chromosome in their search for markers? For that matter, what's a marker?

Each of us carries DNA that is a combination of genes passed from both our mother and father, giving us traits that range from eye color and height to athleticism and disease susceptibility.

One exception is the Y chromosome, which is passed directly from father to son, unchanged, from generation to generation. Unchanged, that is unless a mutation -- a random, naturally occurring, usually harmless change -- occurs.

The mutation, known as a marker, acts as a beacon; it can be mapped through generations because it will be passed down from the man in whom it occurred to his sons, their sons, and every male in his family for thousands of years. In some instances there may be more than one mutational event that defines a particular branch on the tree.

This means that any of these markers can be used to determine your particular haplogroup, since every individual who has one of these markers also has the others.

When geneticists identify such a marker, they try to figure out when it first occurred, and in which geographic region of the world. Each marker is essentially the beginning of a new lineage on the family tree of the human race.

Tracking the lineages provides a picture of how small tribes of modern humans in Africa tens of thousands of years ago diversified and spread to populate the world. A haplogroup is defined by a series of markers that are shared by other men who carry the same random mutations.

The markers trace the path your ancestors took as they moved out of Africa. It's difficult to know how many men worldwide belong to any particular haplogroup, or even how many haplogroups there are, because scientists simply don't have enough data yet.

One of the goals of the five-year Genographic Project is to build a large enough database of anthropological genetic data to answer some of these questions. To achieve this, project team members are traveling to all corners of the world to collect more than 100,000 DNA samples from indigenous populations.

The Camacho Ancestral Journey

M168: The EarliestCamacho Ancestor
Fast Facts:
• Time of Emergence: Roughly 50,000 years ago
• Place of Origin: Africa
• Climate: Temporary retreat of Ice Age; Africa moves from drought to warmer temperatures and moister conditions
• Estimated Number of Homo sapiens: Approximately 10,000
• Tools and Skills: Stone tools; earliest evidence of art and advanced conceptual skills

Skeletal and archaeological evidence suggest that anatomically modern humans evolved in Africa around 200,000 years ago, and began moving out of Africa to colonize the rest of the world around 60,000 years ago.

The man who gave rise to the first genetic marker in the Camacho lineage probably lived in northeast Africa in the region of the Rift Valley, perhaps in present-day Ethiopia, Kenya, or Tanzania, some 31,000 to 79,000 years ago. Scientists put the most likely date for when he lived at around 50,000 years ago.

His descendants became the only lineage to survive outside of Africa, making him the common ancestor of every non-African man living today.

But why would man have first ventured out of the familiar African hunting grounds and into unexplored lands? It is likely that a fluctuation in climate may have provided the impetus for the Camacho ancestors' exodus out of Africa.

The African ice age was characterized by drought rather than by cold. It was around 50,000 years ago that the ice sheets of northern Europe began to melt, introducing a period of warmer temperatures and moister climate in Africa. Parts of the inhospitable Sahara briefly became habitable.

As the drought-ridden desert changed to a savanna, the animals hunted by these ancestors expanded their range and began moving through the newly emerging green corridor of grasslands. These nomadic ancestors followed the good weather and the animals they hunted, although the exact route they followed remains to be determined.

In addition to a favorable change in climate, around this same time there was a great leap forward in modern humans' intellectual capacity.

Many scientists believe that the emergence of language gave us a huge advantage over other early human species. Improved tools and weapons, the ability to plan ahead and cooperate with one another, and an increased capacity to exploit resources in ways we hadn't been able to earlier, all allowed modern humans to rapidly migrate to new territories, exploit new resources, and replace other hominids.

M89: Moving Through the Middle East
Fast Facts:
• Time of Emergence: 45,000 years ago
• Place: Northern Africa or the Middle East
• Climate: Middle East: Semiarid grass plains
• Estimated Number of Homo sapiens: Tens of thousands
• Tools and Skills: Stone, ivory, wood tools

The next male ancestor in the Camacho ancestral lineage is the man who gave rise to M89, a marker found in 90 to 95 percent of all non-Africans. This man was born around 45,000 years ago in northern Africa or the Middle East.

The first people to leave Africa likely followed a coastal route that eventually ended in Australia. These ancestors followed the expanding grasslands and plentiful game to the Middle East and beyond, and were part of the second great wave of migration out of Africa.

Beginning about 40,000 years ago, the climate shifted once again and became colder and more arid. Drought hit Africa and the grasslands reverted to desert, and for the next 20,000 years, the Saharan Gateway was effectively closed. With the desert impassable, these Camacho ancestors had two options: remain in the Middle East, or move on. Retreat back to the home continent was not an option.

While many of the descendants of M89 remained in the Middle East, others continued to follow the great herds of buffalo, antelope, woolly mammoths, and other game through what is now modern-day Iran to the vast steppes of Central Asia.

These semiarid grass-covered plains formed an ancient "superhighway" stretching from eastern France to Korea. These ancestors, having migrated north out of Africa into the Middle East, then traveled both east and west along this Central Asian superhighway. A smaller group continued moving north from the Middle East to Anatolia and the Balkans, trading familiar grasslands for forests and high country.

M9: The Eurasian Clan Spreads Wide and Far
Fast Facts:
• Time of Emergence: 40,000 years ago
• Place: Iran or southern Central Asia
• Estimated Number of Homo sapiens: Tens of thousands
• Tools and Skills: Upper Paleolithic

The next Camacho ancestor, a man born around 40,000 years ago in Iran or southern Central Asia, gave rise to a genetic marker known as M9, which marked a new lineage diverging from the M89 Middle Eastern Clan.

His descendants spent the next 30,000 years populating much of the planet. This large lineage, known as the Eurasian Clan, dispersed gradually over thousands of years. Seasoned hunters followed the herds ever eastward, along the vast super highway of Eurasian steppe. Eventually their path was blocked by the massive mountain ranges of south Central Asia—the Hindu Kush, the Tian Shan, and the Himalayas.

The three mountain ranges meet in a region known as the "Pamir Knot," located in present-day Tajikistan. Here the tribes of hunters split into two groups. Some moved north into Central Asia, others moved south into what is now Pakistan and the Indian subcontinent. These different migration routes through the Pamir Knot region gave rise to separate lineages.

Most people native to the Northern Hemisphere trace their roots to the Eurasian Clan. Nearly all North Americans and East Asians are descended from the man described above, as are most Europeans and many Indians.

M45: The Journey Through Central Asia
Fast Facts:
• Time of Emergence: 35,000 years ago
• Place of Origin: Central Asia
• Climate: Glaciers expanding over much of Europe
• Estimated Number of Homo sapiens: Approximately 100,000
• Tools and Skills: Upper Paleolithic

The next marker of the Camacho genetic heritage, M45, arose around 35,000 years ago, in a man born in Central Asia. He was part of the M9 Eurasian Clan that had moved to the north of the mountainous Hindu Kush and onto the game-rich steppes of present-day Kazakhstan, Uzbekistan, and southern Siberia.

Although big game was plentiful, the environment on the Eurasian steppes became increasing hostile as the glaciers of the Ice Age began to expand once again. The reduction in rainfall may have induced desert like conditions on the southern steppes, forcing these ancestors to follow the herds of game north.

To exist in such harsh conditions, they learned to build portable animal-skin shelters and to create weaponry and hunting techniques that would prove successful against the much larger animals they encountered in the colder climates.

They compensated for the lack of stone they traditionally used to make weapons by developing smaller points and blades -- microliths -- that could be mounted to bone or wood handles and used effectively.

Their tool kit also included bone needles for sewing animal-skin clothing that would both keep them warm and allow them the range of movement needed to hunt the reindeer and mammoth that kept them fed.

These ancestors' resourcefulness and ability to adapt was critical to survival during the last ice age in Siberia, a region where no other hominid species is known to have lived.

The M45 Central Asian Clan gave rise to many more; the man who was its source is the common ancestor of most Europeans and nearly all Native American men.

M207: Leaving Central Asia
Fast Facts:
• Time of Emergence: 30,000 years ago
• Place of Origin: Central Asia
• Climate: Glaciers expanding over much of Europe and western Eurasia
• Estimated Number of Homo sapiens: Approximately 100,000
• Tools and Skills: Upper Paleolithic

After spending considerable time in Central Asia, refining skills to survive in harsh new conditions and exploit new resources, a group from the Central Asian Clan began to head west towards the European subcontinent.

An individual in this clan carried the new M207 mutation on his Y chromosome. His descendants ultimately split into two distinct groups, with one continuing onto the European subcontinent, and the other group turning south and eventually making it as far as India.

The Camacho lineage falls within the first haplogroup, R1, and gave rise to the first modern humans to move into Europe and eventually colonize the continent.

M173: Colonizing Europe—The First Modern Europeans
Fast Facts:
• Time of Emergence: Around 30,000 years ago
• Place: Central Asia
• Climate: Ice Age
• Estimated Number of Homo sapiens: Approximately 100,000
• Tools and Skills: Upper Paleolithic

As these ancestors continued to move west, a man born around 30,000 years ago in Central Asia gave rise to a lineage defined by the genetic marker M173. His descendants were part of the first large wave of humans to reach Europe.

During this period, the Eurasian steppelands extended from present-day Germany, and possibly France, to Korea and China. The climate fostered a land rich in resources and opened a window into Europe.

The Camacho ancestors' arrival in Europe heralded the end of the era of the Neandertals, a hominid species that inhabited Europe and parts of western Asia from about 29,000 to 230,000 years ago. Better communication skills, weapons, and resourcefulness probably enabled my ancestors to outcompete Neandertals for scarce resources.

This wave of migration into Western Europe marked the appearance and spread of what archaeologists call the Aurignacian culture.

The culture is distinguished by significant innovations in methods of manufacturing tools, more standardization of tools, and a broader set of tool types, such as end-scrapers for preparing animal skins and tools for woodworking.

In addition to stone, the first modern humans to reach Europe used bone, ivory, antler, and shells as part of their tool kit. Bracelets and pendants made of shells, teeth, ivory, and carved bone appear at many sites. Jewelry, often an indication of status, suggests a more complex social organization was beginning to develop.

The large number of archaeological sites found in Europe from around 30,000 years ago indicates that there was an increase in population size.

Around 20,000 years ago, the climate window shut again, and expanding ice sheets forced these ancestors to move south to Spain, Italy, and the Balkans. As the ice retreated and temperatures became warmer, beginning about 12,000 years ago, many descendants of M173 moved north again to repopulate places that had become inhospitable during the Ice Age.

Not surprisingly, today the number of descendants of the man who gave rise to marker M173 remains very high in Western Europe. It is particularly concentrated in northern France and the British Isles where it was carried by ancestors who had weathered the Ice Age in Spain.

M343: Direct Descendants of Cro-Magnon
Fast Facts:
• Time of Emergence: Around 30,000 years ago
• Place of Origin: Western Europe
• Climate: Ice sheets continuing to creep down Northern Europe
• Estimated Number of Homo sapiens:
• Tools and Skills: Upper Paleolithic

Around 30,000 years ago, a descendant of the clan making its way into Europe gave rise to marker M343, the defining marker of the Camacho R1b haplogroup. The Camachos are a direct descendent of the people who dominated the human expansion into Europe, the Cro-Magnon. The Cro-Magnon are responsible for the famous cave paintings found in southern France.

These spectacular paintings provide archaeological evidence that there was a sudden blossoming of artistic skills as these Cro-Magnon ancestors moved into Europe. Prior to this, artistic endeavors were mostly comprised of jewelry made of shell, bone, and ivory; primitive musical instruments; and stone carvings.

The cave paintings of the Cro-Magnon depict animals like bison, deer, rhinoceroses, and horses, and natural events important to Paleolithic life such as spring molting, hunting, and pregnancy. The paintings are far more intricate, detailed, and colorful than anything seen prior to this period.

These Camacho ancestors knew how to make woven clothing using the natural fibers of plants, and had relatively advanced tools of stone, bone, and ivory. Their jewelry, carvings, and intricate, colorful cave paintings bear witness to the Cro-Magnons' advanced culture during the last glacial age.

This is where Camacho genetic trail, as is known today, ends. However, as additional data are collected and analyzed, more will be learned about the history of the men and women who first populated the Earth.

-----------
Distribution of R1a (purple) and R1b (red) in Europe, Asia and North Africa
Pamphlet from National Geographic: "The Genographc Project: National Geographic Maps the Human Family Tree" -- front and back.

Notes: More information on the Genographic Project (2005-2010)

https://www3.nationalgeographic.com/genographic/. (All the information on this page was provided by genebase. The Genographic Map on the right, detailing R1a and the Camacho-ancestor-haplogroup-R1b (in red) proportions of today's population, is from Wikipedia.

It is noteworthy that scientists were suprised that all of humanity was rooted in Africa (they had originally assumed Asia). After it was known that Africa was the "cradle," scientists were next surprised that most of Europe was populated by persons who moved from Africa to Asia to Europe and not from Africa directly following the coastine of the Mediterranean. There is still debate over the origins of Modern Humans: Multiregional or Out of Africa (2001 article).
http://eternidad.tribalpages.com/
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1 comentario:

  1. Anónimo4 de febrero de 2012, 15:06

    Dear Maria Cecilia, I'm so glad I found you. I am a Native New Yorker of Colombian descent. Like you, and many other Colombians, and Latin Americans of Galician, Asturian and Basque descent, I became curious to find out more about my heritage. I remember my grandfather as a little girl telling me his family came from the land of the Godos. I was little and I thought Godos were mythical creatures. I lost my grandparents and the wealth of knowledge that went with them. As I got older I realized Godos was the spanish word for Goth. I traveled to Spain in my youth to town where my great grandfathre was from. It was quite a journey where I soon discovered that it was a celtic land with celtic customs. And it was no wonder that I had always been mistaken for Irish by all my Irish American friends here. I then read the stories of the Lebor Gabala about King Breoghan and Milesius who conquered Ireland. I realized this had to be more than mythology, and must be proven somehow. I came across works by Professor Sykes of Oxford. And I came upon a comment you made on a message board stating you were a Colombian living in London who had already undergone genetic testing. I began to search for you. I am so extremely excited to find yout his way. And to find that you are a wealth of knowledge on the subject of genetics. I lived in Spain in the Basque region for many years, and I realized all these Basque surnames were also very much concentrated in parts of Colombia in Antioquia, a very insular part of Colombia for centuries. The people are so fair, their spirit is so unique. I remember my grandmother called them Vasconcelos. With such unique names as Echeverry, Irizarry, Aranguren which I soon realized were names with meanings in the Basque language. I realized there has to be proof of our common ancestry with the inhabitants of the British Isles. And I just read your blog with complete fascination to find that all of my suspicions were true, and had been proved. That you so much Cecilia for this wonderful blog. If you like, I can be reached at lgaliciantree@aol.com

    ResponderEliminar

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Latin genes

Latin America is the home of dances as varied as the stately rumba and the racy salsa. It ranges over searing deserts, rocky heights, and sultry jungles. Perhaps it should not be a surprise, as a new study published this week in the Proceedings of the National Academy of Sciences shows, that its people run the genetic gamut as well.

Researchers from NYU and Stanford gathered DNA from 100 people, each with ancestry from one of four Latin American populations: Colombian, Dominican, Ecuadorian, and Puerto Rican. History tells us to expect that these Latino populations should be a mixture of Native American, European, and African populations. By comparing the Latino DNA with reference individuals’ DNA from these three source populations, using an analysis similar to that of 23andMe’s Ancestry Painting feature, the researchers found stark genetic differences between the Latino groups, and between individuals within each group.

The study found that Latino populations differ markedly in the contribution of the source populations. In Ecuador and Colombia, there is typically little African contribution to ancestry, and most individuals are genetic mixes of European and Native American. But in the Caribbean locations of Puerto Rico and the Dominican Republic, at the western reach of the African slave trade, people typically include substantial African ancestry.

Within each of these populations, individuals vary widely in their genetic ancestry composition. In the Colombian sample, for instance, there are individuals that are essentially fully European, people that are 50/50 European/Native American, and people that are 40/40/20 European/African/Native American.

Besides being a fascinating exploration of natural history, this study suggests that there may be practical consequences to Latin American genetic diversity. This is because medical genetic research can be more difficult in diverse populations. In the case of association studies, the name of the game is to study two groups of people who differ only with respect to whether they have the disease of interest or not; people in the disease group should have the same mix of age, gender, ancestry, etc. as in the non-disease group. The genetic diversity of Latin American populations may hinder such studies because of the difficulty in building genetically-matched groups. The study’s authors suggest that new techniques may need to be developed to carry out genetic research in Latin America.

R1b


R1b1b2a1 is almost unique of West Europe

[Typo: in the maps M529, also known as L21, is wrongly written as M259. My apologies]

This is one of the virtues of Myres' paper (that I mentioned yesterday): that a somewhat more clear phylogenetic subdivision is made, emphasizing the difference between West European R1b1b2a and other R1b or R1b1b2, often blurred in previous papers, causing great confusion even to researchers themselves.

A defect is that instead of using a standard name for the defining SNP (L51/S167 per ISOGG) they chose to name it M412. However Argiedude says it is the same marker and I imagine it is. [Update: confirmed: the rs number indicates it is the same SNP].

Another virtue is that some of the substructure of R1b1b2a1 is also mapped, what really covers pretty well the Northern and Italian area of spread of this lineage and even some relatively unmapped areas of SW Europe, specially France.

In any case the apparent structure is curious, so I got the supplementary table S4 (supp. material is freely accessible) and made this map:

Click to enlarge

Notice that, following Argiedude, M412 stands for L51 and M529 stands for L21, what, if confirmed would make the following equivalences:
  • R1b1b2a-M412 = R1b1b2a1 (L51)
  • R1b1b2a1a2-M529 = R1b1b2a1a2f (L21)
Even if not confirmed, the equivalence should be approximate anyhow.

I'm sorry for the horrible color palette but it's my first attempt to make pie charts with Open Office spreadsheet gadget. Next time I'll do better I hope.

Notice also that I did not use all the samples, in the cases of small countries or less relevant regions I arbitrarily chose and discarded some.

Finally notice that pie charts represent only apportions of R1b1b2 and say nothing of the frequency of the lineage overall, which in most of East Europe and West Asia (excepting Turkey and a few neighbors) is extremely low.

The apparent structure of R1b1b2a1

The most apparent structure is, as we already knew the rather different R1b1b2a1a1 and R1b1b2a1a2 distribution. The first one (color coded brown and light blue) is dominant in the North and rather rare in the South - hence: 'North clade' for short hereafter. The second one (dark green, light green, purple and light orange) is by comparison not just more frequent in the South but also probably more diverse as well - hence: 'South clade'. However it is also found in the North.

Then there are some transitional "remnants": R1b1b2a1a* (L11) and R1b1b2a1* (M412). These should be informative (meaning some extra diversity at their structural levels) in order to infer the history of the haplogroup.

Per the hierarchical distribution seen here and diversity data from older works, the most likely origin of R1b1b2 as a whole is Anatolia.

Then R1b1b2a1* (M412) (yellow) is suggestive of a Mid-Danubian (or Italian or Iberian) coalescence.

R1b1b2a1a* (L11) (middle green) is suggestive of a West-Central European (or SW European) coalescence. More data on the Pyrenean region would clarify this maybe.

And, after this layer, comes the division into the widespread Southern and Northern clades mentioned before.

A reasonable interpretation is that the lineage traveled relatively fast upstream of the Danube (and/or via North Italy onto the SW), branching out then into the two major North/South clades. These two lower level lineages are in fact the two main stars of this demographic expansion.

My bet is that this represents a wave of colonization of Europe (when?) with secondary expansions from SW Europe (Franco-Cantabrian region possibly) and Central Europe (Rhine-Danube region I presume). There are several scenarios that can account for this, essentially Paleolithic (or pushing into the Epipaleolithic).

I don't see clearly how this structure could account for a Neolithic spread, really: no Mediterranean Neolithic pattern is apparent at all and Danubian limited expansion cannot account for any spread to SW Europe, certainly not South of the Loire and certainly not at the frequencies it is found there (nor in Britain/Ireland either). Claiming a Neolithic spread of R1b1b2 with this structure can only be done from a very shallow understanding of Neolithic archaeology and prehistory overall.

The main known demographic expansion we know of in the European Upper Paleolithic is the one after the Last Glacial Maximum, when Magdalenian culture expanded from the Franco-Cantabrian region, both northwards to Central Europe and, later, southwards into Iberia. This led to cultural divergence into the Epipaleolithic, with the important expansion into the newly available areas of the Far North, earlier covered in ice. Within the Epipaleolithic some further cultural flows are detected: from the Franco-Cantabrian region into Iberia (Azilian) and from somewhere in Mid-West Europe into the Southwest (Sauveterre-Tardenoisian).

Also, back to the LGM, Magdalenian techno-culture may have got a NW European ultimate inspirational origin but anyhow mediated by the warmer and richer Franco-Cantabrian region, where the culture flourished properly.

It's difficult to reconstruct in detail but, as far as I can tell, the two main North/South clades must have expanded in the Magdalenian period (one from the Franco-Cantabrian region, the other from Central-NW Europe itself) and also in the ulterior Epipaleolithic. Neolithic does not seem able to account for much but may have helped to shake the board a bit, specially in East-to-West direction.


Frequency maps

Selected frequency maps from the paper
Click to enlarge

Notice that the expansion of the South clade to the Atlantic islands does not invalidate its southern character and probably represents an Epipaleolithic-to-Neolithic spread.

Notice also the large amount of unclassified Southern clade in Iberia. The area around the Pyrenees was not really sampled in this study and therefore it is distorted by neighbors ("South France", looking more like SE France, Valencia and Cantabria).

In order to appreciate better the real thing in this aspect it's probably good to take a look at Cruciani 2010, who did bother to sample near the Pyrenees and gets maybe better (or at least complementary) maps illustrating the same problem.


Update: I superimposed (with complementary colors) the South (red) and North (blue) clades from the frequency maps above. However in order to account for the differences of frequency, I had to lighten the blue shade (North clade) because the scales are different. Take it as an "artist's impression" anyhow:



Update (Aug 27):

Here there is a hopefully better version of the map at the beginning of this post:

Click to enlarge

I put special care in giving each distinct clade an specific color range for easier visualization. All R1b1b2a1 (M412/L51/S167) seems to have coalesced in the Central-to-Western European area but the real expansion seems to have happened after this haplogroup split in two, which I dubbed the North and South clades.

And this is my reconstruction of the haplogroup expansion:

Click to enlarge
Color coded as above


Update (Aug 28):

Take a peek at the comment section, where I briefly discuss molecular clock difficulties and also the only possible Neolithic scenario for R1b1b2a1a2 (South clade): a massive demographic expansion in the context of Megalithism.

Rejecting or confirming this would require greater research in the structure hidden "under the asterisk" in SW Europe. At the moment only two minimally-sized sub-haplogroups are known: Basque/Gascon-specific R1b1b2a1a2b and sub-Pyrenean R1b1b2a1a2c (Gascon, Catalan, etc.). This alone gives highest structural diversity to the Pyrenean region, however most of the South clade remains unresolved (hidden under the asterisk), both in the Pyrenean area as in Iberia proper. And the key issue to solve would be if R1b1b2a1a2 is most diverse at the Pyrenees, what favors a Paleolithic spread scenario, or in West Iberia (and Brittany/West France), what would favor a Neolithic-Megalithic spread scenario instead.

Also it's maybe important to remind here the excellent STR work of Laura Morelli earlier this year, which was discussed in this article.

Importantly, this graph (annotated by me):


The graph is suggestive of the existence of another "West Asian" distinct haplogroup "under the asterisk" (that I labeled "R1b1b2a2?") and a possible Balcanic, rather than Anatolian origin for the R1b1b2 clade.

If so, this would correlate with the high diversity of the (much smaller) brother haplogroup R1b1a in the Italy-West Asia arch (as well as in Central Africa) and would suggest a slightly different origin and scatter for R1b as a whole (ref 1, ref 2).

The R haplogoup tree

The following table shows the R-Tree and the various SNPs that have discovered, their location in the tree and where each clade is often found in the world.

Upper half of R-Haplogroup Tree Lower half of R-Haplogroup Tree

Celtic empire

1200BC Start of the Bronze Age Urnfield Culture in central Europe.
1000-750BC Proto-Celtic people of the Urnfield culture dominate much of Continental Europe. Also start to spread out over northern Asia as far as the frontiers of China. Development of the deliberate smelting of iron in the Middle East and China around the same time. Prompting the title 'The Iron Age' for this period.
700-500 Hallstatt culture developes in Austria.
700BC Early Celts in Austria bury iron swords with thier dead.
600BC Greeks found the colony of Massilia, opening up trade between the Celts of inland Europe and the Mediterranean. First evidence of Britain having a name - Albion - (albino, white - called after the chalk-cliffs of Dover). A major rebuild of old Bronze Age defences, and construction of new hillforts takes place in Britain.
550-500 A princess in Vix (Burgundy) is buried with a 280 gallon bronze Greek vase, the largest ever made. 60 miles away a prince is buried layed out on bronze chais-lounge in a hugh chamber tomb.
500 Trade between the Etruscans and the Celts begins. Lá Téne phase of Celtic culture speads through Europe and into mainland Britain. The Greeks record the name of a major tribe - The KELTOI - and this becomes the common name for all of the tribes.
500 Celts (the Gaels - from Galicia) arrive in Ireland from Spain.
400-100BC La Tene culture spreads over Europe and into the British Isles.
400 Celts invade Italy and Cisalpine Gaul.
400 Celts atack the Etruscan city of Clusium.
390 Raiding Celtic tribes under the leadership of Brennus ravage Rome and occupy the city for three months. Offended by the dirty conditions of the city (they were country boys at heart) they demand a ransome to leave the Romans alone. Brennus demands his weight in gold and when the Romans complain he throws his sword on the scales to be weighed as well with the cry "VAE VICTUS" - (Woe to the Vanquished).
335 Alexander recieves envoys from the Celts, and exchange pledges of alliance. Large numbers of Celtic Warriors join the Greeks in a war against the Etruscans.
323 Alexander dies and the Celts push into Macedonia.
279 Celtic tribes invade Greece.
275 Celts establish the state of Galatia (Gauls across the alps) in northern Turkey.
250
230 Galatian Celts defeated in battle by Greek forces in Western Turkey.
225 Roman army routs invading Celtic Gauls at Telamon in central Italy.
200 The Celts establish permanent fortified settlements (Oppida, or towns).
191 Cisalpine Gaul is taken by the Romans.
150
121 Rome takes Provence.
100 Belgae tribes migrate to Britain to escape Roman domination.
70 Druids (a fire cult from the Middle East) arrive in Britain and gain control of the ruling classes.
58 Julius Caesar is made governor of Provence
58-51 Caesar's Gallic Wars
58 Helvettii in Switzerland are attacked by Germanic tribes led by Ariovistus and move to Gaul. Caesar follows them and defeats them at Toulon-sur-Arroux. Dumnorix of the Aedui tries to lead resistance against the Romans and fails.
57 Caesar then turned his attention to the tribes of the Belgae and lays seige to their territory. By autumn, Caesar claims that all the Gallic tribes are subjects of Rome.
56 The Veneti of Brittany seize two Roman envoys, and make a stand. After a long sea battle, Caesar executed the leaders and sold the men of the tribe into slavery.
55 Julius Caesar tries to land in Britain and is pinned down on a beachhead for two months. His cavalry was seasick and was sent back to Gaul. With the aproaching Autumn gales he withdraws from Britain.
54 Caesar prepares another expedition to Britain and attempts to take Dumnorix as a hostage. Dumnorix refuses and the Romans kill him. As he dies he cries "I am a freeman in a free state". Inspired by his actions, Ambiorix of the Eburones leads an attack against the Roman garrison and massacres them. Ambiorix recruits the Belgic tribes, the Nervii and Aduatuci, and lay seige to the garrison at Namur. The attack is so successful that Caesar himself had to lead the relieving army to drive them off.
53 The tribes of Gaul unite under the leadership of Indutiomarus of the Treveri. The Celtic army consisted of the Treveri, Senones, Carnutes, Nervii, Aduatuci and Eburones. Indutiomarus attacks Caesar's headquarters at Mouzon and lays seige. After a great fight, the Romans kill Indutiomarus. There then followed a number of uprisings among the tribes and Caesar has to work his way through the tribes putting down revolts. Acco of the Senones and the Carnutes was flogged and then put to death. Ambiorix was trailed by a Roman troop until he disappeared into the Ardennes forest, and was never heard from again.
52 A war leader called Vercingetorix ( « Read more about him) emerges to take control of the Celtic army. He maintains a running battle from three successive hill forts. The last one was called Aelisia and Ceasar laid siege for three months with no effect and had to defend himself from from constant attack by the Celtic warriors. Vercingetorix finely surrenders.
45 Caesar ordered that Vercingetorix was to be taken to Rome. He was paraded through the streets then executed as a dangerous enemy of Rome.

0

 Birth of Christ. (Acording to the church of Rome under Constantine)
AD38 Caligula parades Celtic captives through Rome.
AD39 The Catevaulauni under the Kingship of Cunobelinus and his sons Caratacus and Togidubnus, expand into the Atrebate (Hampshire) and the Trinovante (Suffolk).
AD41 Petition put in to Rome for assistance, turned down because of the civil wars in Rome.
AD43 Verica of the Atrebates petitions Claudius to come to Britain to help against the Catevaulauni.
AD 43 Claudian invasion with four legions under Aulus Plautius. Defeat of Caratacus and capture of Camulodunum. Expansion into the midlands (XX Valeria Victrix and XIV Gemina) and in the east (IX Hispana). Frontier established west of Fosse Way. Caractacus escapes into the Welsh borders and fights the Romans using guerilla tactics. Once it is safe to do so, Claudius comes to Britain in person to claim it for Rome. He rides an elephant into the new town of Londinium, stays for two weeks, then goes back home.

47

 New governor, Ostorius Scapula, governor, draws a frontier from the Trent to the Severn. Campaigns in the west (Legio II Augusta under Vespasian)

49

 Colonia of Camulodonum (Colchester) founded. And Roman expansion starts into Wales
49- 50 Foundation of Colonia Victricensis at Camulodunum. Mendip lead mines already in Roman hands. Legionary fortresses at Glevum and Lindum. Invasion of South Wales.

50

 Caratacus, finally defeated in North Wales, flees to Cartamandua, queen of the Brigantes, and is surrendered to the Romans.

52

 New Governor, D. Gallus.

c.55

 Didius Gallus, governor, intervenes on the side of Cartimandua in Brigantian civil war.

57

 New Governor, Q. Veranius

58

 New Governor, S. Paulinus, attack on N. Wales.

59-60

 Suetonius clears Britain totally of the Druids, with a final stand on Anglsea.

60

 Suetonius Paulinus, governor, attacks Anglesey.

60

 Pratagustus of the Iceni dies, and the Romans take his lands away from Boudica.

60-61

 Boudica is elected war leader and leads a revolt agianst the Romans. (Read the story) Icenian revolt under Boudica suppressed after sack of Camulodunum, Londinium and Verulamium

63

 New Governor, T. Maximus.

65

 Preparations for campaigns in Wales.

66

 One legion (XIV Gemina) withdrawn from Britain.
68 Army in Britain refuses to join the governor, Trebellius Maximus, in revolt against Galba.
69 Romans fail to prevent the defection of the Brigantes.
69 Civil Wars, New Governor, V. Bolanus.
71 New Governor, P. Cerialis. Conquest of Brigantia, capture of Stanwick?
71-74 Petilius Cerealis, governor, with a new legion (II Adiutrix) conquers the Brigantes. Legionary fortress at Eburacum.
74-78 Sextus Julius Frontinus, governor, subdues Wales and plants garrisons there. Legionary fortresses at Isca and Deva.
78 Julius Agricola, governor, completes the conquest of North Wales and Anglesey.
79 Consolidation of Brigantian conquest.
80 Advance to the Central Lowlands.
81 Agricola advances to the Forth Clyde line.
82 Conquest of south-west Scotland.
83-84 Agricola advances north and defeats the Caledonians at the battle of Mons Graupius. Roman fleet circumnavigates Britain. Legionary fortress at Inchtuthil.
84 After the Battle of Mons Graupius, occupation of N.Scotland.
84-85 Agricola recalled by Domitian.
86 One legion (II Adiutrix) withdrawn from Britain.
c.90 Legionary fortress at Inchtuthil evacuated.
90-96 Foundation of Lindum Colonia at Lincoln.
96-98 Foundation of Colonia Nervia Glevensis at Gloucester.
99-100 Legionary fortress at Isca and many auxiliary forts in Wales rebuilt in stone. Scottish forts evacuated.
c103 Legionary fortress at Deva rebuilt in stone.
107-108 Legionary fortress at Eburacum rebuilt in stone.
117 Revolt in north Britain.
122 Hadrian visits Britain. Legio IX Hispana replaced by VI Victrix. Construction of Hadrian's Wall from Tyne to Solway begun by Aulus Platorius Nepos.
139-142 Q. Lollius Urbicus, governor under Antoninus Pius, advances into Scotland and builds the Antonine Wall across the Clyde-Forth isthmus.
155-158 Rebellion in north Britain suppressed by C. Julius Verus. Antonine Wall temporarily evacuated.
161-165 Forts rebuilt by Calpurnius Agricola.
180-184 Further revolt in north Britain subdued by Ulpius Marcellus. Antonine Wall broken.
193 On the assassination of Commodus, Pertinax (lately governor of Britain) is chosen emperor by the Praetorian Guard but quickly killed. Empire auctioned to Didius Julianus, who is defeated by Severus.
196-197 Clodius Albinus, governor, takes troops from Britain to fight for the throne and is defeated by Severus. Hadrian's Wall, the fortress at Eburacum and many forts over run and destroyed by the Maeatae.
197 Virius Lupus restores the situation and rebuilds many forts.
200-208 Rebuilding of Hadrian's Wall by Alfenus Senecio.
208 Severus, Geld and Caracalla arrive in Britain and prepare for northern campaigning.
209 Severus and Caracalla campaign in Scotland and receive the surrender of the Caledonians.
210 Revolt of the Maeatae and second Scottish campaign.
211 Severus dies at York. Withdrawal to Hadrian's Walland organization of southern Scotland as a protectorate.
212 Caracalla extends Roman citizenship to all free provincials. Britain divided into two provinces.
259-214 Britain a part of the Gallic Empire of Postumus and his successors.
275-287 Saxon pirates in the Channel.
287 Commander of the British fleet, usurps the title of Emperor in Britain and Carausius, northern Gaul and is temporarily recognised by Diocletian and Maximian.
293 Carausius' continental possessions. Caesar reconquers Constantius.
294 Carausius murdered by Allectus, who succeeds him.
296 Britain restored to the legitimate emperors by Constantius, who crosses the Channel and defeats and kills Allectus. Barbarian inroads in the north. Hadrian's Wall and legionary fortresses at Eburacum and Deva rebuilt. Diocletian's reorganisation divides Britain into four provinces,separates the military from the civil administration and institutes new military offices.
306 Constantius, now emperor, with his son Constantine campaigns in Caledonia. Death of Constantius at Eburacum.
313 Edict of Milan grants toleration to the Christian Church.
314 Three British bishops attend the Council of Aries.
343 Constantine visits Britain and pacifies the Caledonian tribes.
360 Julian sends Lupicinus to repel raids of Picts and Scots.
364 Picts, Scots, Attacotti and Saxons raiding Britain.
367 Great invasion of Picts, Scots and Attacotti, aided by Saxon pirates and a simultaneous attack on Gaul by Franks.Treachery in the Wall garrison. Nectaridus, Count of the Saxon Shore, killed and Fullofaudes, Duke of Britain, routed.
369 Count Theodosius, sent by Valentinian I, clears Britain of invaders and restores the Wall. Signal stations built on Yorkshire coast.
383 Magnus Maximus, a military commander in Britain, revolts and conquers Gaul and Spain from Gratian. Hadrian's Wall swamped by invaders and not rebuilt.
388 Maximus defeated at Aquileia by Theodosius.
395 Stilicho improves the defences of Britain.
406 Constantine III, a usurper, strips Britain of troops for his conquest of Gaul and Spain.
410 Honorius tells the civitates of Britain to arrange for their own safety. Quote"...look to your own defences..."
c.446 Last appeal of the British civitates to Aetius.
From this point on - whether you like it or not - Celtic Britain ends, or to be more precise Romano-British rule is no more. The Celtic peoples had spent 400 years mixing and marrying with the Romans and all the other peoples that came in smaller numbers from the Empire. For the last 100 years the Saxons had settled the south of England with the Roman forces unable to stop them. In fact the Romans paid the Saxons to keep the peace. When the Empire collapsed and the troops that remained were recalled, they had to run the gauntlet of the Saxon warriors all the way down to the ships on the coast. Many of them were robbed of their possessions on the way through, and some did not make it at all!

What was left of the Romano-British in the south of Britain rallied around a military leader for a few years and kept the Saxons at bay. This shadowy figure is what all the Arthurian stories are based on.