Ancient DNA in Europe - Jodrell Bank Observatorymcdonald/genetics/report-2017-ancient-revised.pdf · Ancient DNA results 3 40000-8000 BC ... and analysis of 5,000-year-old remains

Contents

Part 2: Ancient DNA and migrations around the U106 origin 2Introduction to ancient DNA testing 2Ancient DNA results 3 40000-8000 BC 3 8000-6000 BC 4 6000-5250 BC 5 5250-4500 BC 6 4500-3400 BC 7 3400-2925 BC 8 2925-2550 BC 9 2550-2140 BC 10 2140-1950 BC 11 1950-1500 BC 12

Ancient DNA in EuropeReport to the R-U106 group

November 2017 editionAuthor: Iain McDonald

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tccttacgtagacactaactttaagacagacagtgaagta

ccgggatgcaccgacgtccacttaactgggtagccccctc

ctacttcagagccccgatccgcaaagcaacgagggtttac

ttttatcccgtagcatttggatggcaatgagtcaccttag

cttcccccgaaggtcgtctacctgctgtgaacgaaggtgc

gcagatctccgtcaccccctggttgctggggtccgtggcg

gcaactcctctccaagggcgcgatgtccgactgcggggaa

ctagctatgagcagactacgtgcggtttaaattaaaaatc

gatgcatactgcccgtgcatcccctcgacatccgagtctg

gagattagatggttgaggggtaagtatggagataaacacg

ggaaaactgtcttgaagtctgagtcggatcggagctatta

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ccgccactagtgagcaggcacttcgtaacattcttcgagc

tatagccatgctaatccgctgaggatttctgtatcattta

tgtgcgatgttggattactaattggaattgtaggcagcct

acgcatcacgactattaagatagcgcaagtgtacatctag

acagctcggtatcttagggttgaaatggccacgggcatgt

taaccgatgttacgcaagccgtcacaggacgcgtaatgta

gcgatccatcggggagcaaacatggtggtcgcgagaggtt

cctctggggcaaaggtgtctcttctagacccggcgtccgg

ttcttgtatagaaggtcaatcgcctaatcgcgaggtagag

ccggcgacattggatacacttagcttacgcaccggtcgcc

ASSOCIATING ANCIENT DNA WITH ARCHEAOLOGICALCULTURES

What we get from ancient DNA is a sequence of called variantspresent in the genes of a single man. Normally, there is enoughinformation to tie that man to a particular culture that lived at thattime. Today, an entire culture cannot be adequately represented by asingle person, or a single family. There is no particular reason to thinkthat it was any different in pre-historic times. Any ancient culture istherefore going to be a mixture of haplogroups. However, each culture will have a different admixture ofhaplogroups, so tracking the ancient migrations of our ancestorsrequires tracking how these haplogroup admixtures change overgeographical space and over time. Over ancient timescales, this is acase of looking at the ratio of haplogroup I to G, I to R, G to R, etc.In more modern times, we need to look at the ratio of morecontemporary haplogroups, like R-U106 to R-P312, etc. The mostimportant thing, where possible, is to look at the spread ofcontemporaneous haplogroups, to maximise the fraction of thathaplogroup within that culture.

The sparse sampling of ancient DNA means that we do not geta complete picture of a haplogroup's distribution. For example, if aculture is sampled by a single haplogroup I individual, that doesn'tmean the rest of the culture was haplogroup I. Similarly, if there arethree or four R-P312 results from a culture, but no R-U106 results,that doesn't mean that R-U106 is not there, merely that it is likely tobe present at a much lower percentage. It's also important toremember that ancient DNA often only samples burials of the eliteclasses (there are many exceptions). This may mean that we onlysample a few ruling families, while the bulk of ancient peoples gounrecorded because they didn't warrant a high-status burial (e.g. alarge tumulus), where their remains would be well-preserved andeasily identifiable. Sequencing of results from battlefields, etc., canhelp rectify this, but there will remain a class bias in our results,particularly in (post-)Neolithic cultures.

As with all datasets, our knowledge improves as more ancientDNA is sequenced. True sequencing of ancient DNA has only reallybeen possible in the last handful of years, so we are only starting toget a real picture of how different haplogroups populated Europe.This is a fast-evolving field, with potentially game-changingdiscoveries being made every few months..

DNA extracted from archaeological remains are an invaluable sourceof the distribution of haplogroups in ancient and prehistoric times,and in recording the clades that had formed by those peoples deaths.They provide a method to validate the ages of nodes in the haplotree,and they provide information we can use to deduce the migrations ofpeople in these ancient times.

This document focusses on ancient DNA results in Europebetween until 1500 BC, after which few useful samples are available.These cover a wide range of periods, from the first modern humansto inhabit Europe, to the arrival of R-L11 into Europe (circa 3000BC), and the spread of R-L11 that broadly defines the currenthaplogroup distributions today. The maps on the following pagesshow these ancient DNA results, broken up by period, to highlight thedifferences between them.

The ancient DNA records used to provide the following mapscome from Jean Manco's database:

http://www.ancestraljourneys.org/ancientdna.shtml

PROBLEMS WITH ANCIENT DNA

There are many difficulties in testing ancient DNA, related to boththe preservation of the sample and possible contamination since thatperson's death, including by the archaeologists and scientistshandling the remains. At the time of writing (November 2017),extracting and sequencing DNA from some 45,000-year-old remainshas been achieved, and analysis of 5,000-year-old remains isbecoming routine. However, the quality of the results is still muchlower than commercial tests and depends strongly on the state ofpreservation. Typically, a modern sequencing test might reach a fewdozen good-quality reads on the same base pair. If you find the sameallele (A,C,G,T) at that base pair (nearly) every time, it gives youconfidence that you have the correct one. With ancient DNA, you arelucky to get one or two reads, though calls are usually only madewhen a higher number of good-quality reads is reached (say 5x).

This means that the effective coverage of ancient DNA is veryvariable, and doesn't cover every SNP that a standard sequencing testwould. Only in a small proportion of samples is the DNA ofsufficiently good quality to call novel variants from the test, and oftenonly broad haplogroups (e.g. "R1b") are published. Several dedicatedindividuals process the publicly-available raw results from these teststo extract further known SNPs. These results often remain tentative,and may still only identify a test as, e.g. R1b-M269, rather than somemore recent variant, but we have to do the best with what we have. The problems associated with ancient DNA are summarised inthe review article by Kivisild (2017):

https://link.springer.com/article/10.1007/s00439-017-1773-z

ANCIENT U106 RESULTS

While this document is intended to be fairly general, it is made forthe R-U106 group. There have only been two confirmed U106+burials discovered in time period we consider here (up to 1500 BC).The first is RISE98, who was found buried in the southernmost tip ofSweden, and appears to have lived at some point during the lattercenturies of the third millennium BC, coming from the SwedishBattle Axe culture, seemingly a descendant culture of the westernCorded Ware Culture. The second is at De Tuithoorn, north ofAmsterdam, and appears to represent a transition population betweenthe subsiding Bell Beaker Culture and a culture of the Nordvestblock. This lack of ancient R-U106 is partly expected. Most ancientburials in nothern, western and central Europe around 3000-2000 BCare R-P312. The top of the R-P312 tree branches much faster thanR-U106. indicating a faster population spread. However, this doesn'tentirely account for the lack of observed R-U106. It seems likely thatR-U106 exists in a region that is not well sampled during the period2500-2000 BC. It may be holed up somewhere like the southernBaltic coast, where few testers have been sampled.

After 2000 BC, the only currently (November 2017) confirmedU106+ burials are from a Roman-era cemetery in York, one of whichis R-DF98 and one of which is R-DF96. It is likely that these burialsrepresent gladiators from within Roman Britain. The isotopicanalysis of the R-DF96 burial suggests that he came from a moremountainous region. The paper's authors posit Wales, but other areasof the British Isles (e.g. southern Scotland) remain possible. Only oneof the burials mentioned in that paper shows isotopic and autosomalDNA evidence from coming from outside the British Isles.

Ancient DNA

40000-8000 BC: Ice Age EuropeUpdated: 12 May 2017

Dr. Iain McDonaldon behalf of the U106/S21 group

R1b-M343R1b1-P25R1b1a-P297

R1b1a2-M269R1b1a2a-L23

R1b1a2a1-L51

R1b1a2a1a-P310

R1b1a2a1a1-U106P312

M478

S1194

Z2118

S1161

Z2103

L277.1

L584Z2106

PF7558

221

13

6

2

10

2

10

10

122 5

2600 BC (3050 BC - 2100 BC)

2700 BC (3450 BC - 2150 BC)

3500 BC (4450 BC - 2800 BC)

4000 BC (5000 BC - 3100 BC)

4100 BC (5350 BC - 3200 BC)

The dark grey area shows the time periodnominally covered by each chart. Thelight grey area shows the same period,allowing for uncertainties in dates. (Forthis first chart, they are identical.)

11300 BC (12900 BC - 9700 BC)

KEYHaplogroup I:Haplogroup G:Other non-R:Haplogroup R1a:Haplogroup R1b:R1b-P311:R1b-P312:R1b-U106:Dark symbols: carbon-14 or context dateswhere the average of the 95% confidenceinterval dates overlaps with the stated agerange. Lighter symbols: burials where the95% c.i. partially overlaps with the statedrange, but the central estimator does not.

KostenkiC1b-M130; Fu+2016; 36730-34310 BCCT; Fu+2016; 31040-29890 BC

GoyetC1a2; 33210-32480 BCHIJK; 13280-12830 BCFu+2016

Dolni VestoniceCTxIJK, BT; 29120-28720 BCC1a2, F; 28760-27360 BCPosth/Fu+2016;

VillabrunaR1b1a-L754Fu+2016*12230-11830 BCEpigravettian

BichonI2a1a2aJones+201511820-11610 BCAzalian

Ust-IshimK2aPosnik+201647480-42560 BCNot included in count

PeteracuOaseK2aPosnik+201639690-35530 BC

Grotta PaglicciI-CTS674Fu+201632630-29260 BC

PeteraCioclovinaUscatCTPosth/Fu+201631140-29830 BC

PavlovIJK

Fu+201629160-27460 BC

Hohle FelsI-CTS674Fu+201614050-12310 BCMagdalenian

BurkhardtshhleI-CTS5650Fu+201613130-12200 BCMagdalenian

SatsurbliaJ1

Fu+2016*11430-11180 BC

RochedaneI-CTS4088Olalde+1711140-10880 BCEpipaleolithic

IboussiresOlalde+1710090-9460 BCEpipaleolithicJ?-P209R-F356

PadinaR1b1a(x1a)-A702; 9221-8548 BCI2a1-P37.2; 8753-8351 BCR1b1a(x1a)-A702; 8703-8246 BCR1b1a(x1a)-A702; 9500-6200 BCOlande+2017

VasilevkaR1a-L62/L63; 8825-8561 BCI2a2a-P220; 8280-7967 BCOlande+2017*Mesolithic

Boncuklu2*G2a2b2b

Kilinc+20168300-7952 BC8212-7952 BC

VlasacI-CTS7502Olande+20178240-7940 BC

Composition9 (36%)012 (64%)1 (4%)3 (12%)000Typical uncertainties: 20%. Non-Rpredominantly CT before 13,000 BC andI thereafter. Additional untyped R notincluded. Timescale is pre-M269. R-P297appears by 12,000 BC in south-eastEurope.

Appr

oxim

ate lim

it of

glacia

l ice,

20 00

0 BC

No ancient DNA is found in northernEurope during this period due to theextensive ice cover. This reached amaximum around 20 000 years ago, butgradually disappeared as the climatewarmed and sea levels rose. By circa 8000BC, both were close to todays levels.

The earliest remains in Europe belong toa variety of haplogroups not generallyfound in Europe today, including C1a2(C-V20). Some early remains (e.g. GrottaPaglicci) are of extant Europeanhaplogroups, such as I2.

The series of remains from circa 13,000 -8000 BC are dominated by haplogroup I2,especially in the northern regions(although this may be a sampling effect).Also present are an early R1b clade, R-A702 (e.g. Villabruna), which may bemore common in southern regions.

8000-6000 BC: The Neolithic revolutionUpdated: 17 May 2017


R1b-M343R1b1-P25R1b1a-P297


R1b1a2a1-L51

R1b1a2a1a-P310

R1b1a2a1a1-U106P312

M478

S1194

Z2118

S1161

Z2103

L277.1

L584Z2106

PF7558

221

13

6

2

10

2

10

10

122 5

2600 BC (3050 BC - 2100 BC)

2700 BC (3450 BC - 2150 BC)

3500 BC (4450 BC - 2800 BC)

4000 BC (5000 BC - 3100 BC)

4100 BC (5350 BC - 3200 BC)

11300 BC (12900 BC - 9700 BC)


PadinaR1b1a(x1a)-A702Olande+20179500-6200 BC

VasilevkaI2a2a-P220; 8280-7967 BCOlande+2017*Mesolithic

Tepecik CiftlikG2a2a; 6635-6475 BC

2*C1a2-V1526; 6345-6215 BCKilinc+2016

VlasacI-CTS7502; 8240-7940 BCR1b1a(x1a,x2)-A702; 7100-5900 BCI2-PF3664; 6655-6225 BCI-CTS7946; 6500-6250 BCI2a2a-L622; 6200-5900 BCI2a2a1b-CTS10100; 6200-5900 BCI2a2a1b2-Z161; 6200-5900 BCOlande+2017

W/C Eur Fmr. USSR Balkans Anatolia3 (100%) 0 9 (56%) 3 (16%)0 0 1 (6%) 9 (47%)0 1 (33%) 0 7 (37%)0 1 (33%) 0 00 1 (33%) 6 (38%) 00 0 0 00 0 0 00 0 0 0Composition is split by: western/central Europe; former USSR; Balkans &Anatolia. Typical uncertainties: >50%, >50%, 25%, 23%. Non-R predominantly Ibefore 6,300 BC with G2, C1, H2 arriving in south-east afterwards. Additionaluntyped R not included above. Timescale is pre-M269. R-P297 appears commonacross eastern Europe.

Kotias KlbeJ2a

Fu+2016*7940-7600 BC

Ostrovul CorbuluiR1b1a(x1a)-A702; 7580-7190 BCOlande+2017

ZvejniekiR1b1a1a(x2)-CTS3876; 7465-7078 BCOlande+2017

Falkenstein-HhleI2a2a-P223Olande+20177460-7040 BC

Schela CladoveiI2-PF3664; 7340-6640 BCR-M764; 7050-6530 BCOlande+2017

Hajducka VodenicaI2a2-L35; 7300-6000 BCR1b1a-A702(x1a,x2); 7300-6000 BCI2a2a1b2-Z161; 6355-5990 BCOlande+2017

Yuzhnyy Oleni OstrovR1a1a1-M459; 6850-6000 BCMatheson+2015

BarnHykG2a2b2a-CTS688

G2a2b2a3-Z393183*G2a2a1a2a-PF3237

2*C1a2-V20/V15263*H2-L279I-CTS7946

Mathieson+20156500-6200 BC

G2a2b2a1a1c-CTS3426067-5982 BC

ChaudardesI-CTS2387Fu+20166410-6100 BC

DzhulyunitsaG2a2b2b1a-F872;

6300-6150 BCC-F1367;

6061-5841BCOlande+2017

LoschbourI2a1b-L178Lazardis+20146210-5990 BC

Lepenski VirR1b1a-CTS4244; 6222-5912 BCOlande+2017

MenteseG2a-CTS653; G2a2a1-S15710;

I-CTS2387Kilinc+2016

6400-5600 BC

During the early Holocene, the smallnumber of European results indicate thathaplogroup I2 was still dominant incentral Europe. However, lack of reliabledata makes interpretation difficult.

The sparse set of eastern European resultsfrom the early Holocene suggests thathaplogroups R1a and R1b were alreadywell established in this region, during atimeframe corresponding to thepopulation bottleneck between R-P297and R-M269.

Data from the Balkans and Anatoliamarks the interface between theindigenous Mesolithic haplogroups I2and R-A702 and the incoming Neolithicfirst farmers, most of whom arehaplogroup G2a2. Haplogroup C1a2 isstill present in Anatolia, although is notapparent in Europe during this period.

DereivkaI-CTS2387; I2a2a1b-CTS10100; I2a2a1b1-L702R-M734; R-M651; R1-CTS997; R1a-L62; R1b1-PF6250;5*R1b1a(3*x1a,1a2)-A702/CTS4244/PF62495500-4800 BCOlande+2017(3 relatives ignored)

6500-5250 BC: Farmers in the BalkansUpdated: 17 May 2017


R1b-M343R1b1-P25R1b1a-P297


R1b1a2a1-L51

R1b1a2a1a-P310

R1b1a2a1a1-U106P312

M478

S1194

Z2118

S1161

Z2103

L277.1

L584Z2106

PF7558

221

13

6

2

10

2

10

10

122 5

2600 BC (3050 BC - 2100 BC)

2700 BC (3450 BC - 2150 BC)

3500 BC (4450 BC - 2800 BC)

4000 BC (5000 BC - 3100 BC)

4100 BC (5350 BC - 3200 BC)

11300 BC (12900 BC - 9700 BC)


VlasacI2a2a-L622; 6200-5900 BCI2a2a1b-CTS10100; 6200-5900 BCI2a2a1b2-Z161; 6200-5900 BCOlande+2017(Relative ignored)

W/C Eur Scand. Fmr. USSR Balkans Anatolia2 (40%) 4 (100%) 7 (42%) 2 (12%) 1 (33%)0 0 0 6 (38%) 2 (67%)3 (60%) 0 2 (12%) 8 (50%) 00 0 1 (6%) 0 00 0 7 (42%) 0 00 0 0 0 00 0 0 0 00 0 0 0 0Composition is split by: western/central Europe; Scandinavia; former USSR; Balkans & Anatolia.Typical uncertainties: >40%, >50%, 24%, 24%, >50%. Mixture of mostly I2 and R1b1a in Europe withG2, C1, H2 confined to Balkans. G2, C1 spreads thereafter. Additional untyped R not included above.Timescale is pre-M269. R-P297 appears common across eastern Europe, with unclear contribution overwestern and central Europe.

Hajducka VodenicaI2a2a1b2-Z1616355-5990 BCOlande+2017

BarnHykG2a2b2a1a1c-CTS342

Mathieson+20156067-5982 BC

DzhulyunitsaC-F1367

6061-5841 BCOlande+2017

LoschbourI2a1b-L178Lazardis+20146210-5990 BC

Lepenski VirR1b1a-CTS42446222-5912 BCOlande+2017

MenteseG2a-CTS653; G2a2a1-S15710;

I-CTS2387Kilinc+2016

6400-5600 BC

YabalkovoG2a2a1a2a-PF3238; 6000-5900 BC

I2a2-L35; 5726-5575 BCOlande+2017

La Brana-ArinteroC1a2-V20

5983-5747 BCMatheson+2015

GovrlevoG2a2b2b1-F1193

Olande+20175979-5735 BC

MotalaI2a1b-M423; 5964-5638 BCI2a1-P37.2; 5964-5629 BCI2a1b2a1-L147.2; 5721-5631 BCI2c-FGC18140; 5714-5575 BCMathieson+2015; Olande+2017

Beli Manastir-Popova zemlja

C-CTS31515837-5659 BCOlande+2017

ZvejniekiR1b1a1a-CTS3876; 5841-5636 BC4*R1b1a1a(x2)-PF6475& 2*I2a1-P37.2; 2*I2a2a1-CTS9183;2*I2a2a1b-CTS10057; Q1a2-CTS26566000-5100 BCR1b1a1a(x2)-PF6475; 5302-4852 BCOlande+2017

Tiszaszolos-DomahazaI2a1-P37.2; 5780-5640 BCMathieson+2015

CotatcuG2a2b2b-CTS5434

Olande+20175715-5626 BC

Alsonyek-BataszekH2-L281

5702-5536 BCMathieson+2015

StarcevoMalak PreslavetsT1a1-PF5658; 5800-5400 BCC-CTS6266; 5800-5400 BC2*G2a2b2a-CTS10366; 5800-5400 BCOlande+2017(Relative ignored)

LebyanzhinkaR1b1a1a-CTS38765657-5541 BCMathieson+2015

Zemunica CaveC1a2-V20

E1b1b1a1b1-CTS32875600-5470 BCOlande+2017

GomolavaG2a2a1-PF3148

5604-5376 BCOlande+2017

VovnigiI2a2a1b1b-L6995473-5326 BCOlande+2017

Berry au bacI-CTS25145370-5220 BCOlande+2017

VolnienskyI2a2-L35; I-FGC2412;I-FGC24156500-4000 BCIJ-M429; I-CTS8333I2-M438; I2a2a-L59I2a2a1b1-L7025500-4800 BCOlande+2017(IJ interpreted as I)(1 relative ignored)

Yuzhnyy Oleni OstrovJ-CTS59345500-5600 BCMathieson+2015

Els TrocsR1b1a(x1a2)-A702; 5294-5066 BCI2a1b1-L161;5308/5310-5080/5078 BCMathieson+2015

KarsdorfT1a-M70; 5500-4850 BCMathieson+2015LBK

SchletzC1a2-M1009

5500-4775 BCOlande+2017

LBKKleinhadersdorf

Flur MarchleitenJ2-L228

5500-4775 BCOlande+2017

LBK

Kompolt-KigyoserC1a2-V205295-4950 BCMathieson+2015LBK

This period sees I2 remainingcommonplace in parts of western andcentral Europe, but samples are sparse.

The Balkans and south-easternEurope sees an influx of Neolithichaplogroups during this period. C1a2is commonplace, marking a possiblerevival for this haplogroup in Europe.The R-A702 haplogroup is stillpresent, but represents a small part ofthe overall European population,even in the south. Haplogroup Gmakes more of an incursion intoEurope as well.

In the east, there is a mixture of I2,R1a and R1b haplogroups. While thetotal number of burials is significant,the total number of sites are few,raising the possibility of relatednessamong the individuals. Compared tolater populations, R1b-A702 appearscommon, and R1a much rarer.

5250-4500 BC: Farmers in EuropeUpdated: 23 May 2017


R1b-M343R1b1-P25R1b1a-P297


R1b1a2a1-L51

R1b1a2a1a-P310

R1b1a2a1a1-U106P312

M478

S1194

Z2118

S1161

Z2103

L277.1

L584Z2106

PF7558

221

13

6

2

10

2

10

10

122 5

2600 BC (3050 BC - 2100 BC)

2700 BC (3450 BC - 2150 BC)

3500 BC (4450 BC - 2800 BC)

4000 BC (5000 BC - 3100 BC)

4100 BC (5350 BC - 3200 BC)

11300 BC (12900 BC - 9700 BC)


W Europe C Europe Fmr. USSR Balkans3 (75%) 0 11 (42%) 1 (10%)0 5 (56%) 0 4 (40%)0 4 (44%) 1 (4%) 3 (30%)0 0 2 (8%) 01 (25%) 0 9 (35%) 1 (10%)0 0 0 00 0 0 00 0 0 0Composition is split by: western and central Europe; former USSR; and Balkans.Typical uncertainties: >50%, 33%, 20%, 33%. Mixture of mostly I2 and R1b1a before5,100 BC with G2, C1, H2 confined to Balkans. G2, C1 spreads thereafter, C generallyleads G. Additional untyped R included but not shown above. Timescale is probablypre-M269. R-P297 appears common across eastern Europe and into Spain, but coverageacross western/central Europe unclear.

ZvejniekiR1b1a1a-CTS3876; 5841-5636 BC4*R1b1a1a(x2)-PF6475& 2*I2a1-P37.2; 2*I2a2a1-CTS9183;2*I2a2a1b-CTS10057; Q1a2-CTS26566000-5100 BCR1b1a1a(x2)-PF6475; 5302-4852 BCOlande+2017

Berry au bacI-CTS25145370-5220 BCOlande+2017

VolnienskyI2a2-L35; I-FGC2412;I-FGC24156500-4000 BCIJ-M429; I-CTS8333I2-M438; I2a2a-L59I2a2a1b1-L7025500-4800 BCOlande+2017(IJ interpreted as I)(1 relative ignored)

Els TrocsR1b1a(x1a2)-A702; 5294-5066 BCI2a1b1-L161;5308/5310-5080/5078 BCMathieson+2015

KarsdorfT1a-M70; 5500-4850 BCCT-CTS1217; 5217-5041 BCCT; 5216-5036 BCMathieson+2015LBK

ZvejniekiI-CTS2387; I2a2a1b-CTS10100; I2a2a1b1-L702R-M734; R-M651; R1-CTS997; R1a-L62; R1b1-PF6250;5*R1b1a(3*x1a,1a2)-A702/CTS4244/PF62495500-4800 BCOlande+2017(3 relatives ignored)

SchletzC1a2-M1009

5500-4775 BCOlande+2017

LBK

KleinhadersdorfFlur Marchleiten

J2-L2285500-4775 BCOlande+2017

LBK

Kompolt-KigyoserC1a2-V205295-4950 BCMathieson+2015LBK

Halberstadt-Sonntagsfeld4*G2a2a(3*a1)

Various: 5212..5201-5009..4850 BC

Lipson/Olande+2017LBK

SchletzG2a2b2a3-Z39318

5500-4500 BCOlande+2017

LBK

Beli Manastir-Popova zemljaI-FI24790-4558 BCOlande+2017

IvanovoG2a2b2a1a1c1a

-CTS44724725-4605 BC

Mathieson+2017Copper Age

Clos de Roque2*I2a1b4710-4540 BC4680-4460 BCOlande+2017

Gomolava2*G2a2a1a

-PF31774710-4504 BC4605-4460 BCOlande+2017

Neolithic

Apc-BerekalyaC1a2-V205211-4992 BCMathieson+2015LBK

KhvalynskR1a1(1*M459)R1b1a-A702Q1a-F26765200-4000 BCMathieson+2015

VarnaG2-CTS1900; 4685-4499 BCCT; 4685-4406 BCR1-M306; 4551-4374 BCMathieson+2017Copper AgeSmyadovoR-P280R1b1a-A7024550/4545-4455/4450 BCMathieson+2017

Western populations now see the fullinflux of Neolithic farmers, whoappear to reach central Europe, butwhose presence is not heavily felt inwestern Europe or the British Isles.C1a2-V20 is still common in mid-eastern Europe.

Easternmost European populationsstill dont see a large influx ofNeolithic haplogroups, remainingmostly I2 and R1b-A702. Some R1aand other haplogroups are present,but not at modern percentages.

4500-3400 BC: The spread of copperUpdated: 24 May 2017


R1b-M343R1b1-P25R1b1a-P297


R1b1a2a1-L51

R1b1a2a1a-P310

R1b1a2a1a1-U106P312

M478

S1194

Z2118

S1161

Z2103

L277.1

L584Z2106

PF7558

221

13

6

2

10

2

10

10

122 5

2600 BC (3050 BC - 2100 BC)

2700 BC (3450 BC - 2150 BC)

3500 BC (4450 BC - 2800 BC)

4000 BC (5000 BC - 3100 BC)

4100 BC (5350 BC - 3200 BC)

11300 BC (12900 BC - 9700 BC)


Brit. Isles W/C Europe Fmr. USSR Balkans10 (100%) 3 (60%) 0 3 (60%)0 0 3 (38%) 1 (20%)0 1 (20%) 4 (50%) 2 (40%)0 0 0 00 1 (20%) 1 (13%) 00 0 0 00 0 0 00 0 0 0Composition is split by: British Isles; western/central Europe; former USSR; andBalkans. Typical uncertainties: 32%, >45%, 35%,>45%. Fewer burials during thisperiod, possible archaeologically attested population decline. Timescale may be pre-M269. R-P297 still present in northern continental Europe, but notable absence ofhaplogroup R results overall. Neolithic haplotypes now present in Ukraine. No impactin Scotland.

Clos de Roque2*I2a1b4710-4540 BC4680-4460 BCOlande+2017

Gomolava2*G2a2a1a

-PF31774710-4504 BC4605-4460 BCOlande+2017

Neolithic

KhvalynskR1a1(1*M459)R1b1a-A702Q1a-F26765200-4000 BCMathieson+2015

VarnaG2-CTS1900; 4685-4499 BCCT; 4685-4406 BCR1-M306; 4551-4374 BCMathieson+2017Copper Age

SmyadovoR-P280R1b1a-A7024550/4545-4455/4450 BCMathieson+2017

Apc-BerekalyaI2a1-P37.24491-4357 BCOlande+2017

SushinaCT-M56114450-4264 BCMathieson+2017Copper Age

Areni3*L1a14330-4060 BC4230-4000 BC4350-3500 BCLazardis+2016Copper Age

ZvejniekiR1b1a1a-PF64754251-3976 BCMathieson+2017Middle Neolithic

Verteba CaveE-CTS10894; 4000-3600 BCG2a2b2a-CTS688/PF33303931-3640 BC; 3911-3659 BCG2a-PF3141; 3758-3636 BCG2a2b2a1a1b1a1a1-L2;3619-2936 BCMathieson+2017Trypillia

McArthur CaveI2a2a; 3952-3781 BC

Distillery CaveI2a2a1; 3701-3640 BC

I2a2a1a1a; 3514-3353 BCRaschoille Cave

I; I2a2a; I2a1b1; I2a2a1a1a3800-3200 BCOlande+2017

EsperstedtI-CTS33833970-3910 BCMathieson+2015

AbonyG2a2b2a3910-3650 BC2*I2c3800-3600 BC3760-3635 BCLipson+2017

La MinaH2-L279I2a2a1b2-Z1613900-3600 BCMathieson+2015

UrziceniC1a2a-S239813761-3645 BCMathieson+2017Copper Age

QuedlinburgR1b1a(x1a2)

-CTS42443624-3527 BC

Mathieson+2015Baalberge

Tulloch of AsseryI2a1b1; 3635-3390 BC

Olande+2017

El PortalonI2a2a2

3879-3010 BCGunther+2015

Holm of PapaI2a2a1b; 3500-3360 BC

I2a1b; 3510-3340 BCOlande+2017

BatalonlelleI; I2; I2a1a1a3600-3000 BCLipson+2017

Budakalsz2*G2a2b2a1a(1*1b1)

3600-3000 BCG2a2b2a1a1c1a

3367-3103 BCLipson+2017

Remedello di SottoI-CTS40883483-3107 BCAllentoft+2015

OetziG2a2a1a2a1a

3484-3104 BCKelle+2014

The first significant surveys of the British Isles(although only Scotland is covered) show a populationentirely dominated by haplogroup I2a.

Western European populations during this period showa population of mostly I2a and G2a2, reaching someequilibrium. R1b-A702 may still be present at a lowerlevel (e.g. Quedlinburg), but with a much lowerpercentage than seen during the Mesolithic. C1a2 alsoseems to be losing its presence around this time.

Few results are available from the formerUSSR during this period, which isunfortunate, as R-M269 is expected toform here during this period. The Neolithichaplogroup G2a has spread to westernparts of the Ukraine by this time. The fewother samples suggest a population ofmostly R1a and R1b.

3400-2925 BC: The rise of the YamnayaUpdated: 15 June 2017


R1b-M343R1b1-P25R1b1a-P297


R1b1a2a1-L51

R1b1a2a1a-P310

R1b1a2a1a1-U106P312

M478

S1194

Z2118

S1161

Z2103

L277.1

L584Z2106

PF7558

221

13

6

2

10

2

10

10

122 5

2600 BC (3050 BC - 2100 BC)

2700 BC (3450 BC - 2150 BC)

3500 BC (4450 BC - 2800 BC)

4000 BC (5000 BC - 3100 BC)

4100 BC (5350 BC - 3200 BC)

11300 BC (12900 BC - 9700 BC)


Brit. Isles W/C Europe Fmr. USSR Balkans9 (100%) 12 (63%) 0 4 (80%)0 7 (37%) 1 (14%) 00 0 0 1 (20%)0 0 0 00 1 6 (86%) 00 0 0 00 0 0 00 0 0 0Composition is split by: British Isles; western/central Europe; former USSR; andBalkans. Typical uncertainties: 33%, 23%, >38%, >45%. Timescale includes M269 andZ2105. R-P297 now absent in Europe. Neolithic haplotypes now present in Spain. Noimpact in Scotland.

Verteba CaveG2a2b2a1a1b1a1a1-L2;3619-2936 BCMathieson+2017Trypillia

Raschoille CaveI; I2a2a; I2a1b1; I2a2a1a1a

3800-3200 BCOlande+2017

Tulloch of AsseryI2a1b1; 3635-3390 BC

Olande+2017

El PortalonI2a2a2

3879-3010 BCGunther+2015

Holm of PapaI2a2a1b; 3500-3360 BC

I2a1b; 3510-3340 BCOlande+2017

BatalonlelleI; I2; I2a1a1a: 3600-3000 BCI2; 3337-3024 BCLipson+2017

Budakalsz2*G2a2b2a1a(1*1b1)

3600-3000 BCG2a2b2a1a1c1a

3367-3103 BCLipson+2017


OetziG2a2a1a2a1a

3484-3104 BCKelle+2014

Banbury LaneI2a1b; I2a2; I2a2a1

3360-3100 BCOlande+2017

AlsonemediG2a2a3359-3098 BCLipson+2017

EsperstedtI2a1b1a1-S27033360-3086 BCMathieson+2015

Salzmuende-SchiebzigG2a2a-PF31853400-3025 BCMathieson+2015

DzhulyunitsaH2-L279Mathieson+20173336-3028 BC

SmyadovoI2a2a1b-CTS10100; 3338-3025 BCI2a2a1b1-L702; 3328-3015 BCMathieson+2017

IbisterI2a1b; 3336-3024 BC

I2a1b1; 3336-3012 BCI2a1b1; 3334-2942 BC

I2; 3180-2780 BCI2a2a1a1a2; 3011-2886 BC

I2a2; 3009-2764 BCOlande+2017

KierzkowoI2a2-P218; 3335-3020 BCI2a2a1b-CTS10057; 3400-2800 BCI2-L68; 3135-2935 BC*Mathieson+2017Globular Amphorae+2 relatives (ignored)*Approx. date from son

AlavaI2a2; 3263-2903 BCI2a2a2a; 3350-2750 BCG2a2a; 3092-2918 BCI2a2; 3090-2894 BCLipson+2017

QuoynessI2a1b; 3098-2907 BC

Olande+2017

LopatinoR1b1a1a2a2-Z2105; 3339-2918 BCR1b1a1a2-L1353; 3321-2921 BCR1b1a1a2a-L23; 3300-2700 BCR1b1a1a2a2-CTS1078; 3300-2700 BCMathieson+2015Yamnaya

PeshanyR1b1a1a2-CTS85913334-2635 BCAllentoft+2015Yamnaya

MerichleriI2a2a1b1b-L699; 3020-2895 BCMathieson+2017

MednikarovoI2a2a1b1b-L699; 3012-2900 BCMathieson+2017

GkhemI23050-2750 BCSkoglund+2014

KutulukR1b1a1a2-CTS86653335-2882 BCMathieson+2015Yamnaya

British populations during this period remain resolutelycomprised of I2a. No evidence of incursion byNeolithic haplogroups is apparent, and few suchhaplogroups are present in the British population today,suggesting the Neolithic revolution in Britain wascarried as a cultural package, rather than by movementof people.

Continental European populations are now largelycomprised of I2a and G2a2, which are the maincomponents of these populations among todayspopulations. Haplogroups C1a2 and R-A702 seem tohave essentially died out in Europe by this time.

The growth of the Yamnaya culture during this time isdominated by R1b-Z2105 and other R-M269 lines. Nodata from certified R-L51 individuals themselves isfound during this time, so the R-L51 homeland cannotbe claimed to be identified. However, thecontemporaneous growth with R-Z2105 (from circa3700 BC) suggests that it could also be linked with therise of the Yamnaya.

MerichleriI2a2a1b1b-L699; 3020-2895 BCMathieson+2017

2925-2550 BC: The Corded Ware CultureUpdated: 17 July 2017


R1b-M343R1b1-P25R1b1a-P297


R1b1a2a1-L51

R1b1a2a1a-P310

R1b1a2a1a1-U106P312

M478

S1194

Z2118

S1161

Z2103

L277.1

L584Z2106

PF7558

221

13

6

2

10

2

10

10

122 5

2600 BC (3050 BC - 2100 BC)

2700 BC (3450 BC - 2150 BC)

3500 BC (4450 BC - 2800 BC)

4000 BC (5000 BC - 3100 BC)

4100 BC (5350 BC - 3200 BC)

11300 BC (12900 BC - 9700 BC)

KEYHaplogroup I:Haplogroup G:Other non-R:Haplogroup R1a:Haplogroup R1b:R1b-P311:R1b-P312:R1b-U106:Dark symbols: carbon-14 or context dateswhere the average of the 95% confidenceinterval dates overlaps with the stated agerange. Lighter symbols: burials where the95% c.i. partially overlaps with the statedrange, but the central estimator does not.Burials with 95% c.i. starting after 2571BC and central estimators after 2410 BCare not shown in this case.

Brit. Isles C. Europe S. Europe Fmr. USSR Balkans2 (100%) 1 (17%) 9 (82%) 2 (17%) 00 0 1 (9%) 0 2 (67%)0 0 1 (9%) 0 00 3 (50%) 0 1 (8%) 00 2 (33%) 0 9 (75%) 1 (33%)0 0 0 0 00 0 0 0 00 0 0 0 0Composition is split by: British Isles; central Europe (Denmark, Germany, Poland); southern Europe (Spain, Italy)former USSR (Russia, Ukraine); and Balkans. Typical uncertainties: >70%, >40%, 30%, 29% and >57%. Timescaleincludes M269 and may include L11. R-M269 appears in Europe by:* Poland: 2837-2672 BC (M269 or equivalent but see: http://www.anthrogenica.com/showthread.php?10565/page15);* Croatia: 2884-2666 BC (Z2105);* Germany: possibly 2880-2630 BC (R1); probably 2868-2580 BC (R1b1).

IbisterI2; 3180-2780 BC

I2a2a1a1a2; 3011-2886 BCI2a2; 3009-2764 BC

Olande+2017

AlavaI2a2; 3263-2903 BCI2a2a2a; 3350-2750 BCG2a2a; 3092-2918 BCI2a2; 3090-2894 BCI2a2a; 2916-2714 BCLipson+2017

QuoynessI2a1b; 3098-2907 BC

Olande+2017

LopatinoR1b1a1a2a2-Z2105; 3339-2918 BCR1b1a1a2-L1353; 3321-2921 BCR1b1a1a2a-L23; 3300-2700 BCR1b1a1a2a2-CTS1078; 3300-2700 BCR1b1a1a2a2-CTS1078; 2887-2666 BCMathieson+2015Yamnaya

PeshanyR1b1a1a2-CTS85913334-2635 BCAllentoft+2015Yamnaya

MednikarovoI2a2a1b1b-L699; 3012-2900 BCMathieson+2017

GkhemI23050-2750 BCSkoglund+2014

KutulukR1b1a1a2-CTS8665; 3335-2882 BCR1b1a1a2a2-Z2105; 2867-2486 BCMathieson+2015Yamnaya/Poltavka

EkaterinovkaR1b1a1a2a2-CTS10782921-2762 BCMathieson+2015Yamnaya

LuzkhiR1b1a1a2a2-Z20153021-2635 BCMathieson+2015Yamnaya

DzhulyunitsaG2a2a1a2-L912906-2710 BCMathieson+2017

IlyatkaI2a2a1b-CTS10057; 2899-2706 BCI2a2a1b2-Z161; 2890-2694 BCMathieson+2017Globular amphorae

El MiradorI2a2a2a; 2900-2679 BCI-CTS4209; 2900-2346 BCI-CTS5650; 2900-2346 BCI2a1a1a-L672; 2900-2346 BCG2a2b2b1a1-PF33792900-2346 BCMathieson+2015

El PortalonH2; 2899-2678 BCGunther+2015

CaravacaI; 2900-2670 BCOlande+2017

Beli Manastir-Popova zemljaR1b1a1a2a2-Z21052884-2666 BCMathieson+2017Vucedol

TermtaR1b1a1a2a2-CTS9416; 2887-2634 BCR1b1a1a2-PF6482; 3000-2400 BCR1b1a1a2a2-Z2105; 3000-24000 BCAllentoft+2015YamnayaTiefbrunn

R1-CTS5611; 2880-2630 BCR1b1-L1349; 2868-2580 BCAllentoft+2015Corded Ware

SamborzecR1b1a1a2 [M269]2837-2672 BCOlande+2017Bell BeakerDate uncertain?

Vucedol tellG2a2a1a2a-PF32372872-2617 BCMathieson+2017


GrachevkaR1b1a1a2-L2652872-2583 BCMathieson+2015Yamnaya

KierzkowoI2a2a1b22870-2575 BCMathieson+2017Globular amphorae

Sok RiverR1b1a1a1b2a-F31052925-2491 BCMathieson+2015Poltavka outlier

Stalingrad QuarryR1b1a1a2a2c-Z21062857-2497 BCAllentoft+2015

KyndelseR1a1a1-Page72851-2492 BCAllentoft+2015

CerdanyolaI2a22833-2480 BCOlande+2017

BergrheinfeldR1a1a1-M4172829-2465 BCAllentoft+2015Corded Ware

Cheddar GorgeI; 2830-2461 BCOlande+2017

CaravacaCF; 2920-2340 BCMathieson+2017

KalavanR1b1a12619-2465 BCLazardis+2016

UlanI2a2a1b1b2-S121952849-2143 BCAllentoft+2015

NikolaevkaR1b1a1a2a2-CTS10782800-2200 BCMathieson+2015Poltavka

Beli BreyagI2a2-L37I-L7583400-1600 BCMathieson+2017

VibyR1a1a1-Page72621-2472 BCAllentoft+2015

Populations in the British Isles and Iberia remaincomprised almost exclusively of I2a and G2a2, as inprevious and modern populations. A lack of data inFrance and the Low Countries means we have limitedinformation on some areas, although under the CordedWare model, we expect at least France to be a mix ofI2a and G2a2 as well. This period corresponds to theearly part of the Bell Beaker culture in these areas,confirming that it was initiated by I2a+G2a2-dominated cultures.

Populations in central Europe begin to bereplaced by Corded Ware culture peoples duringthis period. The Corded Ware A horizon(according to Furholt 2014) ranges from the2900s BC in the eastern Baltic, through the 2800sin central Europe, arriving on the North Seashores by the 2700s. Burials during this periodare most commonly R1a1a1-M417. TheTiefbrunn burial is not called below R1b1, butcould be R-L11, suggesting R-L11 may havebeen present during this process too. The route bywhich R-L11 arrived in central Europe remainsuncertain, and could have been anywhere from upthe river Danube, to along the southern Balticcoastline.

Southern European populations remaincomprised of I2a and G2a2 during this time,reflecting the Neolithic population. The presenceof R-Z2105 in Croatia shows that incursion byYamnaya-associated forces had occurred insouth-eastern Europe by 2666 BC. ModernZ2105 percentages indicate that this group didnot spread westwards of the Balkans in greatnumbers.

Eastern groups appear dominated by R-Z2105from the Yamnaya populations. There are someI2a2 burials among them, notably the GlobularAmphorae culture burials of the Ukraine andPoland. Its not clear whether this culture shouldrepresent a migrating indigenous Europeanculture, as Y-DNA would suggest, or a trueYamnaya-based cultural element, as thearchaeology suggests.

El MiradorI-CTS4209; 2900-2346 BCI-CTS5650; 2900-2346 BCI2a1a1a-L672; 2900-2346 BCG2a2b2b1a1-PF33792900-2346 BCI2a2a2a-L1228; 2568-2346 BCMathieson+2015

2550-2140 BC: The Bell Beaker CultureUpdated: 17 July 2017


R1b-M343R1b1-P25R1b1a-P297


R1b1a2a1-L51

R1b1a2a1a-P310

R1b1a2a1a1-U106P312

M478

S1194

Z2118

S1161

Z2103

L277.1

L584Z2106

PF7558

221

13

6

2

10

2

10

10

122 5

2600 BC (3050 BC - 2100 BC)

2700 BC (3450 BC - 2150 BC)

3500 BC (4450 BC - 2800 BC)

4000 BC (5000 BC - 3100 BC)

4100 BC (5350 BC - 3200 BC)

11300 BC (12900 BC - 9700 BC)

KEYHaplogroup I:Haplogroup G:Other non-R:Haplogroup R1a:Haplogroup R1b:R1b-P311:R1b-P312:R1b-U106:Dark symbols: carbon-14 or context dateswhere the average of the 95% confidenceinterval dates overlaps with the stated agerange. Lighter symbols: burials where the95% c.i. partially overlaps with the statedrange, but the central estimator does not.Burials with 95% c.i. ceasing before 2460BC and central estimators before 2645BC, or beginning after 2210 BC are notshown in this case.

Brit. Isles Iberia C. Europe Scandinavia S.E. Europe Fmr USSR2 (22%) 7 (78%) 0 0 4 (44%) 1 (25%)0 1 (11%) 1 (3%) 0 1 (11%) 00 0 0 0 1 (11%) 00 0 8 (24%) 1 (50%) 0 1 (25%)0 1 (11%) 10 (29%) 0 2 (22%) 2 (50%)2 (22%) 0 5 (15%) 0 0 05 (56%) 0 10 (29%) 0 1 (11%) 00 0 0 1 (50%) 0 0Composition is split by: British Isles; Spain/Portugal; western/central Europe from France to Poland, Austria to the Netherlands;Scandinavia (southern Sweden); Hungary and the Balkans; and the former USSR. Typical uncertainties: 33%, 33%, 17%, >50%,33% and >50%. Timescale includes major clades of P312 and U106. R-M269 appears in Europe by:* Poland: 2837-2672 BC (M269 or equivalent);* Croatia: 2884-2666 BC (Z2105);* Germany: possibly 2880-2630 BC (R1); probably 2868-2580 BC (R1b1);* France: 2566-2299 BC (P312);* Hungary: 2465-2205 BC (L2);* England: 2460-2200 BC (P311);* Scotland: 2287-2039 BC (DF13);* Sweden: 2275-2032 BC (U106);* Netherlands: 2278-1914 BC (P312).

TermtaR1b1a1a2-PF6482; 3000-2400 BCR1b1a1a2a2-Z2105; 3000-2400 BCAllentoft+2015Yamnaya

CaravacaCF; 2920-2340 BCR1b1a(x1a2a); 2850-2250 BCI; 2460-2140 BCMathieson+2017

VibyR1a1a1-Page72621-2472 BCAllentoft+2015

Osterhofen-AltenmarktR1b1a1a2a1a2-P3122572-2512 BCR1b1a1a2a1-L512500-2000 BCAllentoft+2015

KalavanR1b1a12619-2465 BCLazardis+2016

IbisterI2a1b1; 2581-2464 BCI2a1b; 2571-2348 BCOlande+2017


NikolaevkaR1b1a1a2a2-CTS10782800-2200 BCMathieson+2015Poltavka

UlanI2a2a1b1b2-S121952849-2143 BCAllentoft+2015

CerdanyolaR1b1; 2571-2350 BCG2; 2474-2300 BCOlande+2017Bell Beaker

AlavaI2; 2571-2347 BCI2a2a; 2481-2212 BCLipson+2017

Les Villas d'Aurele2*R1b1a1a2a1a2 [P312]2566-2299 BC2468-2278 BCOlande+2017

CsajerszkeG2a2a1a2a12559-2301 BCOlande+2017

EsperstedtR1a1a1-M4172559-2296 BCR1a1a; R; R1a;R1a1; R; R1a2500-2050 BCMathieson+2015Corded Ware

Harmanoren-Gondurle HoyukJ1a-CTS5368

Lazardis+20172558-2295 BC

SamborzecR1b1a1a2 [M269]; 2571-2208 BCR; 2463-2142 BCOlande+2017Bell Beaker

AugsburgG2a2a1a2a1a; 2471-2300 BCR1b1a1a2a1a [P311]; 2461-2210 BCR1b1a1a2a1a [P311]; 2800-1800 BCR1b1a1a2a1a2b1 [L2]; 2397-2149 BCR1b1a1a2-L150.1; 2500-2000 BCOlande+2017

VillardR1b1a1a2a1a2b1 [L2]2459-2242 BCOlande+2017

SzigetszentmiklsR1b1a1a2 [M269]; 2500-2200 BCI2a2a; 2459-2206 BCR1b1a1a2a2 [Z2105]; 2458-2202 BCI2a1a1; 2458-2154 BCOlande+2017Bell Beaker

Arroyal 1I2a2a; 2461-2210 BCI2a2a; 2458-2206 BCOlande+2017

Budapest-BksmegyerR1b1a1a2a1a2b1 [L2]

2465-2205 BCH2; 2470-2060 BC

Olande+2017

AmesburyR1b1a1a2a1a [P311]; 2460-2200 BCR1b1a1a2a1a2c [L21]; 2470-2140 BCR1b1a1a2a1a2c [L21]; 2480-2031 BCR1b1a1a2a1a [P311]; 2280-2030 BCOlande+2017

QuedlinburgR1b1a1a2-PF6430; 2467-2142 BCR1b1a1a2a1a2-P312; 2341-2150 BCMathieson+2015Bell Beaker

Lkr. DeggendorfR1b1a; ~2300 BCR1b1a1a; ~2300 BCOlande+2017Bell Beaker

RothenschirmbachR1b1a1a2 [M269]2458-2140 BCMathieson+2015

Sur les BarmesR1b1a1a2a1a [P311]2456-2135 BCOlande+2017

MondelangeR1b1a1a2a1a2 [P312]; 2436-2136 BCR1b1a1a2a1a [P311]; 2400-1900 BCOlande+2017Bell Beaker

Stkr. StraubingR1b1a1a2a1a-P311R1b1a1a2a1a2-P3122*R1b1a1a2a1a2b1-L22300-2150 BCAllentoft+2015

YarntonR1b1a1a2a1a2c1[DF13]2400-2040 BCOlande+2017

Cova da MouraI2a1a1; 2289-2135 BCOlande+2017

UtyevkaR1-CTS29082469-1928 BCMathieson+2015Poltavka

BulanovoR1a-L622298-2045 BCAllentoft+2015

Leki MaleR1b1a-L7542286-2048 BCAllentoft+2015CWC/Unetice

West DeepingR1b1a1a2a1a2c1[DF13]2289-2041 BCOlande+2017

Dryburn BridgeR1b1a1a2a1a2c1 [DF13]2287-2039 BCR1b1a1a2a1a2c [P312]2275-1884 BCOlande+2017

KnezevesR1b1a1a2a1a-P3102279-2033 BCOlande+2017

Lille BeddingeR1b1a1a2a1a1-U1062275-2032 BCAllentoft+2015

WillingtonR1b1a1a2a1a2c1 [DF13]2277-1920 BCOlande+2017

De Tuithoorn, OostwoudR1b1a1a2a1a2 [P312]2300-1900 BC2278-1914 BCOlande+2017

This period sees R-L11/P311 Bell Beaker populationsarrive in Great Britain and probably Ireland too. Theyappear to be mostly or wholly R-DF13, arriving before2200 BC in England and 2039 BC in Scotland. BellBeaker archeological results suggest a most-likelyarrival during the 2200s BC, posing an interestingrelationship with the existing Megalithic cultures atsites like Stonehenge and Orkney.

In France and Iberia, the spread of P312 mirrors themovement of the Bell Beaker culture, showing that theBell Beaker reflux into these areas was carried byincoming cultures who appropriated the Bell Beakerphenomenon. The incoming R-M269 dont seem tohave made a significant impact in Iberia, even as late as2289 BC. Existing R1b1 populations (Caravac andCerdanyola) are probably still R-A702.

Northern European groups nowappear a more balanced R1a/R1bmix. Incursion into Swedenseems to occur before 2472 BC.Apart from the Lille Beddingeresult, R-U106 is surprisinglyabsent. The U106/P312 ratio isexpected to remain small until~1500 BC, but the absence stillsuggests that the basal U106population is not being wellsampled during this period.

Central European groups see aP312 concentration, particularlyof U152>L2, spreading down theupper Danube and Rhone, likelyreaching Italy before 2200 BC.This largely seems to replace theI2a and G2a2 populations duringthis time.

Eastern populations are less wellsampled during this period, butprobably remain mostly R1b-Z2015 dominated, with R1a andI2a populations in the mix.


2140-1950 BC: Late Bell BeakerUpdated: 27 July 2017


R1b-M343R1b1-P25R1b1a-P297


R1b1a2a1-L51

R1b1a2a1a-P310

R1b1a2a1a1-U106P312

M478

S1194

Z2118

S1161

Z2103

L277.1

L584Z2106

PF7558

221

13

6

2

10

2

10

10

122 5

2600 BC (3050 BC - 2100 BC)

2700 BC (3450 BC - 2150 BC)

3500 BC (4450 BC - 2800 BC)

4000 BC (5000 BC - 3100 BC)

4100 BC (5350 BC - 3200 BC)

11300 BC (12900 BC - 9700 BC)

KEYHaplogroup I:Haplogroup G:Other non-R:Haplogroup R1a:Haplogroup R1b:R1b-P311:R1b-P312:R1b-U106:Dark symbols: carbon-14 or context dateswhere the average of the 95% confidenceinterval dates overlaps with the stated agerange. Lighter symbols: burials where the95% c.i. partially overlaps with the statedrange, but the central estimator does not.Burials with central estimators before2300 BC are not shown even if their 95%c.i. overlap.

Osterhofen-AltenmarktR1b1a1a2a1-L512500-2000 BCAllentoft+2015

EsperstedtR1a1a; R; R1a;R1a1; R; R1a2500-2050 BCMathieson+2015Corded Ware

AugsburgR1b1a1a2-L150.12500-2000 BCOlande+2017

Budapest-BksmegyerH2; 2470-2060 BC

Olande+2017

AmesburyR1b1a1a2a1a2c [L21]; 2480-2031 BCR1b1a1a2a1a [P311]; 2280-2030 BCR1b1a1a2a1a2 [P312]; 2210-2030 BCR1b1a1a2a1a [P311]; 2140-1940 BCOlande+2017

MondelangeR1b1a1a2a1a2 [P312]; 2436-2136 BCR1b1a1a2a1a [P311]; 2400-1900 BCOlande+2017Bell Beaker

YarntonR1b1a1a2a1a2c1 [DF13]; 2400-2040 BCR1b1a1a2a1a2c1 [DF13]; 2137-1930 BCOlande+2017Bell Beaker

UtyevkaR1-CTS29082469-1928 BCMathieson+2015Poltavka

BulanovoR1a-L622298-2045 BCAllentoft+2015

Leki MaleR1b1a-L7542286-2048 BCAllentoft+2015CWC/Unetice

West DeepingR1b1a1a2a1a2c1[DF13]2289-2041 BCOlande+2017

Dryburn BridgeR1b1a1a2a1a2c1 [DF13]; 2287-2039 BCR1b1a1a2a1a2 [P312]; 2275-1884 BCOlande+2017

KnezevesR1b1a1a2a1a-P3102279-2033 BCOlande+2017

Lille BeddingeR1b1a1a2a1a1-U1062275-2032 BCAllentoft+2015

WillingtonR1b1a1a2a1a2c1 [DF13]2277-1920 BCOlande+2017

De Tuithoorn, Oostwoud4*R1b1a1a2a1a2 [P312]2300-1900 BC2278-1914 BC2196-1903 BC2193-1887 BCOlande+2017

Trumpington MeadowsR1b1a1a2a1a2c1 [DF13]2204-2029 BCOlande+2017

Ingleby BarnwickI2a2a1a1a; 2200-1970 BCOlande+2017

ParmaR1b1a1a2a1a2 [P312]2200-1930 BCOlande+2017

EulauCT-CTS12542137-1965 BCMathieson+2015Not included intotals.

UtyevkaR1a1a1b-S4412200-1900 BCMathieson+2015Poltavka

ReavershillR1b1a1a2a1a [P311]2135-1951 BCOlande+2017

EsperstedtI2c-L5972134-1939 BCMathieson+2015

Eweford CottagesR1b1a1a2a1a2c [P312]; 2140-1916 BCOlande+2017

Szzhalombatta-FldvrI-CTS7722128-1909 BCAllentoft+2015

StepnoeR1a1a1b-S2242126-1896 BCAllentoft+2015

Rathlin IslandR1b1a1a2a1a2c1g [Z30233]; 2026-1885 BC

R1b1a1a2a1a2c1 [DF13]; 2024-1741 BCCassidy+2016


AbeksI-CTS674; 2010-1776 BCAllentoft+2015

Brit. Isles C. Europe Fmr USSR1 (10%) 3 (37%) 00 0 00 0 00 0 1 (100%)0 0 02 (20%) 0 07 (70%) 5 (63%) 00 0 0Composition is split by: British Isles; western/central Europe; and the former USSR. Typical uncertainties: 32%,>35%, >50%. Timescale is post-U106 and post-P312. R-M269 appears in Europe by:* Poland: 2837-2672 BC (M269 or equivalent);* Croatia: 2884-2666 BC (Z2105);* Germany: possibly 2880-2630 BC (R1); probably 2868-2580 BC (R1b1);* France: 2566-2299 BC (P312);* Hungary: 2465-2205 BC (L2);* England: 2460-2200 BC (P311);* Scotland: 2287-2039 BC (DF13);* Sweden: 2275-2032 BC (U106);* Netherlands: 2278-1914 BC (P312);* Ireland: 2026-1885 BC (Z30233).

British populations are uniquely well-sampled duringthis period, indicating a near-complete populationreplacement from I2a to R-DF13. Results from Franceand the Low Countries indicate a strong R-P312population here too.

Central European populations start toshow a modern R1b/R1a fractionationduring this period. The R1 incursiondoesnt seem to have made muchimpact in Hungary, but has clearlyextended as far as Italy. The smallnumber of sites make the exact extentdifficult to determine.

The few samples from Russia duringthis period seem to show the R-Z2105Yamnaya population being surplantedby R1a individuals.

1950-1500 BC: Emerging European CulturesUpdated: 23 November 2017


R1b-M343R1b1-P25R1b1a-P297


R1b1a2a1-L51

R1b1a2a1a-P310

R1b1a2a1a1-U106P312

M478

S1194

Z2118

S1161

Z2103

L277.1

L584Z2106

PF7558

221

13

6

2

10

2

10

10

122 5

2600 BC (3050 BC - 2100 BC)

2700 BC (3450 BC - 2150 BC)

3500 BC (4450 BC - 2800 BC)

4000 BC (5000 BC - 3100 BC)

4100 BC (5350 BC - 3200 BC)

11300 BC (12900 BC - 9700 BC)

KEYHaplogroup I:Haplogroup G:Other non-R:Haplogroup R1a:Haplogroup R1b:R1b-P311:R1b-P312:R1b-U106:Dark symbols: carbon-14 or context dateswhere the average of the 95% confidenceinterval dates overlaps with the stated agerange. Lighter symbols: burials where the95% c.i. partially overlaps with the statedrange, but the central estimator does not.Burials with central estimators before2100 BC are not shown even if their 95%c.i. overlap. Burials with 95% confidenceintervals restricted to before 1900 BC arenot shown.

Brit. Isles Cont. Europe Fmr. USSR0 3 (38%) 00 1 (13%) 00 1 (13%) 00 1 (13%) 6 (86%)1 (17%) 0 1 (14%)0 1 (13%) 05 (83%) 0 00 1 (13%) 0Composition is split by: British Isles; continental Europe and Scandinavia; and the former USSR. Typicaluncertainties: >40%, >35%, >37%.

Dryburn BridgeR1b1a1a2a1a2 [P312]; 2275-1884 BCOlande+2017

De Tuithoorn, OostwoudR1b1a1a2a1a2 [P312]2193-1887 BCR1b1a1a2a1a1 [U106]1881-1646 BCOlande+2017

StepnoeR1a1a1b-S2242126-1896 BCAllentoft+2015

Rathlin IslandR1b1a1a2a1a2c1g [Z30233]; 2026-1885 BC

R1b1a1a2a1a2c1 [DF13]; 2024-1741 BCR1b1a1a2a1a2c [L21]; 1736-1534 BC

Cassidy+2016


AbeksI-CTS674; 2010-1776 BCAllentoft+2015

Hastings HillR1b1a1a2a1a2c1 [DF13]; 1931-1756 BCOlande+2017

Nerquin GetashenR1b1a1a; 1906-1698 BC

Allentoft+2015

ThanetR1b1a1a2a1a2 [P312]1900-1690 BCOlande+2017

ErdI2a2a1a2a[L1229]2000-1500 BCAllentoft+2015

Szreg, Sziv UtcaG2a2a1a2a-FGC77391866-1819 BCAllentoft+2015

UvarovkaR1a1a1b2-F9921850-1600 BCMathieson+2015

Spiridonovka IIR1a1a1b2a2a-Z21231850-1600 BCMathieson+2015

KairyakaNecropolisR1a1a1b2-F9921750-1625 BCMathieson+2017

Boscombe AirfieldR1b1a1a2a1 [L51]1750-1610 BCOlande+2017

Szzhalombatta-FldvrI2a2a1[CTS9183]1746-1611 BCAllentoft+2015

Great OrmeR1b1a1a2a1a2 [P312]1693-1600 BCOlande+2017

UntermeitingenR1b1a1a2a1a [P310]1691-1519 BCAllentoft+2015

Veliki VanikJ2b2a-L2831700-1500 BCMathieson+2017

Spiridonovka IVR1a1a-L1681850-1200 BCMathieson+2015

BarinovkaR1a1a1b2-F9921850-1200 BCMathieson+2015

NovoselkiR1a1a1b2-F9921850-1200 BCMathieson+2015

Documents

Ancient DNA in Europe - Jodrell Bank Observatorymcdonald/genetics/report-2017-ancient-revised.pdf · Ancient DNA results 3 40000-8000 BC ... and analysis of 5,000-year-old remains