Indepth Look at Haplogroup JT

mtDNA haplogroup JT is the daughter of Pan-Eurasian haplogroup R and one of the primary haplogroups in West Eurasia but can also found in many parts of Africa and Asia. It's a very old lineage and covers a lot of land, and therefore has a lot of phylogenetic diversity and regional diversity. In a previous post(LINK) I used HVR1 data from JT samples to display JT regional diversity. To learn more about JT diversity I collected and analysed over 1,000 JT mito-genomes from Ian Logan's site.

I broke this post into four sections to give a good understanding of what I've learned about JT.
1: My Strategy for Analyzing mito-genomes
2: Description of Samples and Spreadsheets used for Analyse
3: Results
4: Comparison to JT mito-genomes from Ancient DNA


My Strategy for Analyzing mito-genomes: My strategy when analyzing mito-genomes is to look for what I call "Expansion Point" haplogroups. An "Expansion Point" haplogroup is a haplogroup which is popular but doesn't have any popular subclades. The reason I this is my strategy is"Expansion Points" are the youngest haplogroups and can tell us the most about the origins of mtDNA in modern populations. They tell us the most about the origins of human mtDNA because most humans belong to "Expansion Point" haplogroups and very rarely do two humans from the same population have maternal lineages that are related beyond being apart of a popular "Expansion Point" Lineage. There are dozens or hundreds of "Expansion Point" haplogroups which take up some 90% of the mtDNA in human populations. Most are fairly young, less than 20,000 years old. For those of you who are familiar with European Y DNA an example of a Y DNA "Expansion Point" haplogroup is R1b-L151.

Description of Samples and Spreadsheets used for Analyse: Most of the samples were used by Maria Pala et al. 2012. They came from various locations in West Eurasia, Siberia, and South Asia. I also added about 500 JT mito-genomes provided by Ian Logan from Denmark, Italy, Arabia, and Iran.

I fully analysed the relationship between all about 1,000 mito-genomes used by Maria Pala et al. 2012. I found over 60 new haplogroups but none of them are popular(aka "Expansion Point" haplogroups) outside of single populations and so not very helpful to know about.

Here's a spreadsheet of that Analysis: Analysis of 1000 JTs.

With the about 500 JT mito-genomes I added to the ones used by Maria Pala et al. 2012, I was able to gather enough samples to create five regional populations to compare their frequencies in JT subclades. The five populations are: Denmark, Italy, USA, Near East, and Northern West Asia. The USA population consists of Americans who's maternal line is from Europe and probably specifically from Britain or Germany.

Here's the spreadsheet of a comparison of those five regions: Regional Frequencies of JT

Results: As I said before my strategy when analyzing mito-genomes is to find "Expansion Point" haplogroups, so here are the "Expansion Point" JT haplogroups I found. I color them according to which of these regions they are most popular in: Generic West Eurasian, Generic Europe, NorthWest Europe, Italy, Generic West Asia, Iran, Near East.

T
T2a1a, T2a1b1a(T2a1b1a1b), T2b(T2b4, T2b23, T2b31), T2c1(T2c1c, T2c1d1, T2c1d1a, T2c1e, T2c1a), T2e, T2f1a1, T2g1a, T2i, T1a1, T1a2, T1a7, T1a11, T1b(T1b2, T1b3).

J
J1c1(J1c1b, J1c1b1a, J1c1d), J1c2(J1c2b, J1c2o), J1c3, J1c15, J1b1a1, J1b1b(J1b1b1), J1b2, J1b3, J1d(J1d1a), J2a1a1a(J2a1a1a2), J2a2a, J2b1a, J2b1(xJ2b1a)

And that's it. Those are all the "Expansion Point" haplogroups of JT. Most JT falls under these clades. There isn't anymore I can do. All subclades of these haplogroups are so rare and exist in most of West Eurasia that they tell nothing about regional diversity. It's amazing there are still pretty basal JT subclades which show no regional variation. T2c1 and T2a1a in Italy and Yeman aren't distinguishable from each other.

Here are a two very important lessons I learned about West Eurasian JT.
Recent contact across all of West Eurasia
>Rare and young non-"Expansion Point" subclades are found in every part of West Eurasia.
>Regional specific and young "Expansion Point" subclades are found in every part of West Eurasia, it's just they're more popular in some parts.
West Asia vs Europe, with Italy and Turkey/Cyprus as intermediates
>NorthWest Asia/Near East and Italy/NorthWest Europe aren't perfect subpopulations, but each member in these subgroups are by far most similar JT to each other. Italy however has a significant amount of typical West Asian JT haplorgoups; e.g. T1a2, T1b, J1b2, J1d1a and Turkey/Cyprus have a significant amount of typical European JT haplogroups; e.g. J1c, J1b1a1, T1a1.
>The majority of West Asian and European JT split well over 8,000 years ago.

Comparison to JT mito-genomes from Ancient DNA

Below is a list of all of the ancient JT mitogenomes I know of. All of the results are from Europe or European ancestors/immigrants who lived in Asia, except for the single Armenian sample. All of them have typical JT clades for modern Europeans, except the Armenian who had T1a2 which is more typical of Western Asia included Northern West Asia(Where Armenia is) today.

6500-6200 BC, Barcın Turkey J1c11
6500-6200 BC, Barcın Turkey J1c
5311-5218 BC Spain J1c3
5000 BC, LBK culture Germany J1c17
2500-2050 BC Corded Ware Germany J1c2e
2880-2630 BC Spain J1c1
3000 BC Spain J1c1b1
2625-2291 BC Corded Ware Germany J1c1b1a
2298-2045 BC Sintashta Russia J1c1b1a
2000 BC? Germany J1c1b
2500-2050 BC Corded Ware Germany J1c5
2290-2130 BC Bell Beaker Germany J1c5
2851-2492 BC Denmark J1c4
2128-1909 BC Hungary J1c9
2863-2498 BC Corded Ware Germany J1b1a1


2880-2630 BC Spain J2b1a3
2126-1896 BC Sintashta Russia J2b1a2a
1850-1200 BC Timber Grave Russia J2b1a2a
1000 BC Siberia J2b1a
3900-3600 BC Spain J2a1a1
2880-2630 BC Spain J2a1a1
2500-2050 BC Germany J2a2a


6500-6200 BC, Barcın Turkey T2b
6500-6200 BC, Barcın Turkey T2b
5000 BC? LBK culture Germany T2b
5000 BC? LBK culture Germany T2b
5000 BC? LBK culture Germany T2b
5000 BC? LBK culture Germany T2b
5000 BC? LBK culture Germany T2b
3360-3086 BC Germany T2b
2500 BC? Czech Republic T2b
2034-1784 BC Hungary T2b3
2000 BC? Hungary T2b
1866-1619 BC Hungary T2b
1800-1600 BC Timber Grave Russia T2b4
794-547 BC Denmark T2b
5178-5066 BC Spain pre-T2c1d2
5000 BC? LBK culture Germany T2c1d'e'f
5100-4800 BC LBK culture Germany T2c1d1
4000 BC? Germany T2c1d1
3000 BC? Yamnaya T2c1a2
5500-4800 BC LBK culture Germany T2e
5000 BC? LBK culture Germany T2e
3640-3510 BC Germany T2e1
2887-2634 BC Yamnaya T2a1a
1432-1292 BC Sweden T2a1a
2454-2291 BC Corded Ware Germany T2a1b1
2464-2210 BC Germany T2a1b1
1800-1400 BC Andronovo Siberia T2a1b

2570-2471 BC Germany T1a1
1395-1132 BC Sweden T1a1
2000 BC? Hungary T1a1
1850-1200 BC Timber Grave Russia T1a1
1048-855 BC Armenia T1a2

Asia Has Five mtDNA Gene Pools

I have added over 2,000 mtDNA samples from Asia and 1,000 from Europe to my mtDNA database in the last few days. Thanks to HaploGrep I've been able to analysis the data more than twice as fast.

With the new data from Asia, I've learned that there are at least Five mtDNA gene pools in Asia. I've also found geographic diversity in East Asian mtDNA and origins of Western mtDNA in Asia.

In the next few weeks I will 1: Research the mtDNA and genome-wide relationship between ancient and modern Siberians, 2: Do more thorough work on European mtDNA with 1,000s of new samples, 3: Collect 1,000s of mtDNA samples from Africa, and 4: Collect many 1,000s of Mito-genomes from Ian Logan which I can now do very quickly thanks to HaploGrep. So, there's a lot to look forward to on this blog if you're interested in mtDNA.

This spreadsheet shows the five mtDNA gene Pools of Asia and regional haplogroups of East Asian mtDNA.

Regional Asian mtDNA

In total there are at least six mtDNA gene pools in Eurasia. Below is a link to a map of the six mtDNA gene pools of Eurasia along with the list of haplogroups in each gene pool.

Eurasian mtDNA Gene Pools

Here are mtDNA Haplogroup frequencies in Asia: Asia mtDNA Frequencies. I included frequencies of South Asian-specfic haplogroups in South Asians and the frequencies of West Eurasian haplogroups in Asia.

Geographic Diversity in East Asian mtDNA

The East Asian mtDNA gene pool is the geographically the largest in the world. Obviously they can't all have the exact same mtDNA. So, I gathered the frequencies of R9 and C subclades in this spreadsheet to find differences: C, R9. East Asia.

There are noticeable differences in subclade frequencies. The biggest is between Siberia and other East Asians. Siberians have a very high frequency of C4a1, C4a2'3'4, and C5 and all three are near non-existent in other East Asians. Tibet/Nepal also have a decent amount of those C subclades, probably because they live near Siberia.

There's also regional-trends in the frequencies of R9 subclades. The most popular R9 subclade in NorthEast and SouthEast Asia is F1a, in Siberia F1b, and in Tibet/Nepal F1c1a. As with C, Tibet/Nepal have a connection with Siberia because of a high frequency of F1b. There's also a high frequency of F4b, R9c1, and F3b in Taiwan and R9b1a in Burma, which are near non existent in other East Asians.

All other East Asian haplogroups can't easily find haplogroups with low-coverage testing, like H in West Eurasia. So, it was hard to find differences in subclade frequencies in other haplogroups, but I did find some. I wrote those differences down here: Regional Asian mtDNA

M7 has consistent strong presence in NorthEast Asia, a weaker presence in SouthEast Asia, and is pretty much unheard of in all other East Asians. M8a consistently pops up in Siberia and NorthEast Asia, and rarely anywhere else. M9a is absent in SouthEast Asia and consistently pops up in other East Asians. E is non-existent in all East Asians except Tawian where it is pretty popular. I also found many D-subclades that are exclusive to certain countries or regions.

NorthEast Asians(Japan, Korea, China) are fairly similar. Tawian and Siberia are differnt from NorthEast Asia in many ways. There are Palaeilthic-splits in mtDNA and more recent links that they have with NorthEast Asia. Nepal and Tibet have a significant amount of South Asian mtDNA, connections with Siberia, and their own unique bottle-neck lineages. Despite Tibet being in the country of China, it's important to remember they aren't Chinese at all, they were just conquered by Chinese.

Origins of Western mtDNA in Siberia and South Asia

West Eurasian mtDNA in Asia peaks in West Siberia and South Asia at about 30%. Every where else in Asia West Eurasian mtDNA is pretty much non-existent. South Asia and Siberia received their Western mtDNA from very differnt sources. Siberian's Western mtDNA is almost entirely from Eastern Europe and South Asian Western mtDNA has unknown sources.

Siberian's Western mtDNA specifically looks like it comes from Pre-Historic Russia(U4, U5a, U2e, T1a, J1b1a1). They probably have a mixture of Mesolithic Russian and Bronze age Andronovo mtDNA(J1b1a1, T1a, J1c, I, H2a1, H6). The composite of Siberian Western mtDNA, is very similar to Catacomb and Andronovo, especially because of their strangely high frequencies of U4.

Siberians also have a string of typical West Asian subclades of U; U7, U1, and U3. It is strange that they don't have a lot non-U West Asian haplogroups. Maybe there were Ancient West Asians who were mostly U7, U1, and U3 like there were ancient Europeans who were mostly U5, U4, and U2e. I doubt it, but it's possible.

South Asian Western mtDNA is dominated by U2(xU2e) and U7. The U2(xU2e) subclades are rarely found outside of South Asia, so have probably been in South Asia for 10,000s of years and not recent arrivals from West Eurasia. The sources of South Asia's U2 is likely a population closely related to Paleolithic North Eurasians, like Ma'lta boy and Kostinki man. The high amount of U7, like in some Siberians, is very strange. U7 is popular in neighbors of South Asia, like Iranians, but it isn't nearly as popular compared to other Western lineages as U7 is in South Asia. Maybe South Asia's U7 and U2 are from the same source.

Non U7 and U2 South Asian Western mtDNA is a mixture of West Asian-specific and European-specific. West Asian-specific mtDNA in my South Asian data besides U7 is includes HV and R0a. Both are more typical of SouthWest Asia than Iran, but still fairly popular in Iran. It's hard to explain the consistent presence of U5a, U4, and J1b1a1, all typical of Bronze age East Europe, in South Asia if all their Western mtDNA is from neighbors in West Asia.

Loads of New mtDNA from Paleo-Europe

Face from a Forgotten World

Ivory carving of Human face dating about 30,000 years old from an archaeological site in the Czech Republic. mtDNA samples from this site had U5* and U8c.

Posth 2016 published several dozen new fully-sequenced mtDNA results from Stone age Hunter Gatherers of Europe(mostly Germany, Belgium, France, Italy). I've added all the results to my mtDNA DataBase. There's now well over 100 mtDNA samples from Stone age European Hunter Gatherers.

The results Posth 2016 are near 100% under haplogroup U(xK), like the results we've been getting from European Hunter Gatherers for years. The only surprise is M* dating about 30,000 years old in Belgium and France.

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Short summary of the history of European mtDNA, up until 6,000 years ago.

30,000+ years ago Europe was settled by humans carrying mostly mtDNA U(U5, U2, U8, etc), but also M*, N*, and R*. Sometime between 30,000 and 15,000 years ago, U5a'b replaced other forms of U in Europe. Western Europe was dominated by U5b and Eastern Europe by U5a, U4, and U2e. No dramatic changes occurred in European mtDNA, till 8,000-6,000 years ago when humans from Turkey and Caucasus mountains brought JT, R0(inlcu. H), U, N1, X, and W.

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The story of European genetics is fascinating and surprising. Modern Europeans are the result of at least two massive migrations from Asia/Far East Europe and a mixture of at least 4 distinct ancestors. They have very close ties to Middle Easterners and to a lesser extent Native Americans and North Asians. In this narrative U-dominated Paleo-Europeans are the aboriginal Europeans. They were some of the first humans to settle Europe and they lived undisturbed for 10,000s of years.

New Ancient mtDNA Reveal Hunter Gatherer Origins of "ENF" mtDNA

There's a type of ancestry that arrived Europe from West Asia with farming in 7000 BC and takes up the majority of ancestry in modern West Asians. It was coined "ENF" by Eurogenes author Davidski.

"ENF" is short for "Early Neolithic Farmer". The reason it was named this is it was believed this type of ancestry expanded out of West Asia with farming

"ENF" arrived in Europe with a package of new mtDNA haplogroups: R0(inclu. H), JT, K, W, X, and N1. Posters at blogs and forums theorized these mtDNA haplogroups must have originated with Early Neolithic West Asian farmers along with "ENF" ancestry. It was a nice and simple story for the origins of Middle Eastern and to a large extent European mtDNA.

I was against this idea. I argued there's just too much mtDNA diversity in West Asia for them to be descended from the same small population only 10,000 years ago. I argued "ENF" and "ENF mtDNA" existed in hunter gatherer populations all over West Asia for 10,000s of years.

In the last few weeks two papers(Hofmanová 2015, Jones 2015) have proven my argument correct. They document "ENF" ancestry in pre-Neolithic Georgia alongside "ENF mtDNA" in pre-Neolithic Georgia and Greece.

7605–7529 BC Thessaly, Greece: K1c
7288–6771 BC Thessaly, Greece: K1c
11000 BC West Georgia: H13c
7700 BC West Georgia: K3

H13c, K3, and K1c are very rare. As far as I know K3 and H13c have basically only been found in the Caucasus region today. And K1c has only been found in Europe, but is very rare.

In Conclusion
These new DNA results reveal the very complicated  and old Paleolithic origins of "ENF" mtDNA. It did not expand with farming. It had been running around in hunter gatherer populations of West Asia(and Europe?) for 10,000s of years. "ENF" itself is a mixture of at least two distinct ancient Eurasian populations, so "ENF" mtDNA is likely also a composite of mtDNA from earlier populations. We need 1000s of mtDNAs from West Asia ranging 45,000 to 3,000 years to understand its origins and history.

Lots of new ancient mtDNA from Anatolia, Spain, and Samara Russia

The new Mathieson 2015(Eight thousand years of natural selection in Europe) included dozens of new mtDNA results from Neolithic Anatolia, Copper age Spain, and Eneolithic-Bronze age Samara Russia. I added the results to my mtDNA DataBase. The results confirm trends revealed earlier by ancient mtDNA. Below are two mtDNA-related highlights from Mathieson 2015.

The mtDNA of (some of)Europe's First farmers came from Anatolia

26 mtDNA samples were taken from people who lived in farming communities at the western edge of Anatolia in circa 6300 BC. The results don't differ at all from mtDNA results of Europe's first farmers in Central Europe and Spain from 5500-5000 BC. Y DNA and autosomal DNA confirm these Neolithic Anatolians were the ancestors of Early Neolithic Central Europeans and Iberians.

It's important to note T2b, J1c, K1a, N1a1a take up the majority of Neolithic Anatolian and Early Neolithic European mtDNA. T2b and J1c today are considered European-specific and modern Europeans certainly inherited them from Neolithic Anatolia(or near by regions). The form of K1a the Neolithic Anatolians carried was all K1a4 and K1a2, except for one K1a3. All these forms of K1a have been found in Neolithic Europe and are popular in Europe today.

U3, U8b, H5, W1c'i, and X2a-o are minor lineages shared between Neolithic Anatolians and Central Europeans. None of the Neolithic Anatolians had U5 or U4 which exists at around 5% in Early Neolithic Europeans, possibly reflecting their 7% "WHG" admixture reported based on autosomal DNA by Mathieson 2015.

Not all Neolithic mtDNA from Europe is very similar to Neolithic Anatolians. mtDNA data from Neolithic Romanians is slightly differnt. They have a much high frequency of H and lack T2 and N1a1a and all their T was T1 like most T in the Balkans today.

Although Neolithic Anatolian and Central Europeans belonged to many European-specific lineages their mtDNA differed in many ways to modern European mtDNA. Whatever happened to N1a1a? Why is K1a and T2b not so popular today? Why is H so popular today? Autosomal DNA tells us Neolithic Anatolians are direct ancestors of modern Europeans, however there's lots of mystery as to why stark differences exist in mtDNA between the two.

T1a1, I, H6a, H2a1 are Steppe-lineages?

I,  H6, and H2a1 don't appear in ancient European mtDNA till Steppe peoples from Ukraine and Russia migrated into Central Europe in the 3rd millennium BC. T1a existed in Neolithic Europe but was at a higher frequency in Steppe populations and first appears in Germany with Corded Ware after being absent for 2,000 years.

The new mtDNA data from Mathieson 2015 adds to the list of Ancient Steppe people who belong to T1a, I, H6a, and H2a1. Various branches of U5a, U4, and U2e are also quite obviously mostly of Steppe-decent in Europe today. Today the highest frequency of Steppe-related mtDNA is found in Volga Russia, Scandinavia, NorthEast Europe, and the Balkans. Steppe-specific subclades are also found in West Asia, Iberia, and Italy. In the Hindu Kush there's a decently high frequency of U4, U2e, and U5a which might have mostly Steppe origins.

The reason the title says T1a1 not T1a, is high coverage Steppe T1a from Mathieson 2015 is specifically T1a1. There are also several examples of T1a1 in Late Neolithic Europeans who had a lot of Steppe-ancestry. Today T1a1 takes up almost 100% of T1a in Europe and a very small percentage of West Asian T1a. It's an obvious Steppe mtDNA haplogroup.

Part 2: Looking for mtDNA diversity in West Eurasia

Previously I showed that within haplogroups JT, R0(xHV), HV(xH), U(xK), N1 diversity follows a West Asia vs Europe pattern. 75%+ of European/West Asian JT, R0(xHV), HV(xH), U(xK), and N1 falls under European/West Asian-specific clades which separated from each other in the Neolithic or earlier.

In today's post I'll show diversity within haplogroup H, shared haplotypes and deep subclades among West Eurasians in JT and UK, and how Pre-Historic European mtDNA is ancestral to modern European mtDNA. Later I'll edit this post and make it more attractive with some PCAs and graphs.

Sorry, I can't give all the details the data reveals. There isn't enough room. I let the data speak for itself. If you don't understand feel free to post any question. 

H diversity is also West Asia vs Europe

In my first post haplogroup H, the most popular haplogroup in West Eurasia, remained undiversified. So I searched on Google for studies that tested H-subclade SNPs. I found several studies with a lot of useful data on H-subclade frequencies in West Asia and Europe. I'm not assuming H diversity follows a West Asia vs Europe trend before I look at the data the data speaks for itself. Here are the results....

H Frequencies, West Eurasia

The most significant differences are that H1 is usually 15% or more in all of Europe and less than 5% in all of West Asia, H3 also has a noticeable high frequency in all of Europe except in the East Baltic, Sweden, and Karelia. H13 has a slightly high frequency in all of West Asia.

The majority of West Asian H is not  H1 through H7 while 75% of European H is. This means more research is needed on West Asian H to learn what sub-lineages the 50%+ other part of West Asian H falls under. In most H sub clades there aren't any significant trends. I'm not going to split hairs and pretend there are trends where there aren't. 

JT, UK haplotype sharing in West Eurasia

The majority of my data is only of HVR1 coverage. This means most of my data can only be put under old haplogroups and diversity is hard to find. The best one can do to find diversity when his data is only of HVR1 coverage is to look for HVR1-haplotype sharing. 

I went through every single  JT/UK HVR1 haplotype and searched for matches. If a haplotype existed in two more populations I found the frequencies of that haplotype in all  populations. 

Here are the results....

JT haplotypes

UK haplotypes

I'm excited about the results for JT and UK. There lots of sharing and regional trends. I broke up the JT and UK haplotypes according to what region it appears they're most popular. I added Pre-Historic matches to the spreadsheet. There are some interesting ancient/modern matches.

How Pre-Historic European mtDNA is ancestral to modern European mtDNA

Here you can see I updated my haplotype sheet with Pre-Historic data. 

I added the following Pre-Historic European meta-populations to the same analysis I did for modern West Eurasians in my first post. 

Pre-Historic European meta-populations:

Early Neolithic Central Europe, Early Neolithic SouthWest Europe, Middle Neolithic North Europe, Middle Neolithic SouthWest Europe, Neolithic Romania, Bronze age Steppe, and Bronze age Central Europe.

Haplogroup Frequencies(Modern and Pre-Historic)

JT+N1: Pre-Historic      Modern.

U: Pre-Historic      Modern 

No Europeans have majority or very significant West Asian mtDNA. All have majority pre-3000 BC European mtDNA. Neolithic Anatolians(ancestors of Europe's first farmers), Mesolithic Europeans, and an unknown Near Eastern ancestor of Yamnaya are the primary mtDNA ancestors of Europeans.

Almost all Pre-Historic European mtDNA fall under European-specific haplogroups. But none have every European-specfic haplogroup. Looking at Pre Historic European mtDNA you can  see which pre-Historic Europeans are the source for each European-specific haplogroup. J1c, T2b, HV0, HV6-17, H1, H3 look like they decend from Europe's first farmers. U5b looks like it descends from Mesolithic Central and West Europeans. U5a, U4, T1a, J1b1a1 T2a1b look like they descend from the Bronze age Steppe.

Basque vs Danish

•  VS 

In my last post I discovered large differences between West Asian and European mtDNA with low coverage mtDNA data. Within Europe  and most of West Asia it is impossible to find such large differences between differnt regions/ethnic groups with low coverage mtDNA data.

Now, I have been able to find a great amount of mtDNA differences between two European ethnic groups: Danish and Basque, because I have high coverage mtDNA data from both of them.

Here's my comparison analysis of Danish and Basque mtDNA.
Basque, Danish mtDNA H
Basque, Danish Other mtDNA
Basque vs Danish mtDNA Frequencies

All of the Danish data was fully-sequenced but for Basque only their H was fully-sequenced.  So, I wasn't able to find a lot of differences between most non-H in Basque and Danish.

H takes up nearly half of European mtDNA and 90%+ of H remains undefined with low coverage testing. Fully-sequenced Danish and Basque H  reveals European H is far from uniform. Nearly 50% of the H from each appears to have separated from each other 8,000+ years ago.

Most noteworthy is that about 60% of H1/H3 in Basque and Danish probably separated from each other 8,000+ years ago. This is noteworthy because H1/H3 take up 40-50% of pan-European H and are very rare in West Asia. They're probably the most important part of the history of H in Europe.

Outside of H Danish differentiation with Basque reflects their higher amount of "Steppe" ancestry(U5a, U4, T1a, J1b1a, J2a1a1, and I).  Outside of H Basque differentiation with Danish mostly reflects Basque-specific founder effects(U5b1f1a, J1b1a-16368, J1c5c1) and a higher amount of "EEF/WHG" ancestry(K1a, HV0(xV), T2(xT2b, T2c1, T2a1b, T2f, T2e)).