Population genetics | Europe | Ancient DNA

Europe's genetic outliers, and why they stand apart

Most Europeans sit on one smooth genetic map. A few populations fall off it: Sardinians, Basques, Saami, Finns, Roma, Kalmyks. We measured how far each population of Europe is from all the others, and found four different ways to become an outlier. The Italians, often cited as an exception, are not one of them.

73residual distance of the Sardinians when rebuilt from all other Europeans, twelve times the European median. Add Neolithic Sardinians as a source and it falls to 12
39%South Asian ancestry in the Roma of the Balkans, the legacy of a founding group that left India about 1,500 years ago
3.0residual of the Tuscans, as ordinary as the English or the Poles. Italy is a gradient between Europe and the eastern Mediterranean, not an outlier

Hero image: AI generated illustration of a Bronze Age village around a dry-stone nuraghe tower in the hills of Sardinia, around 1400 BC, when the islanders still carried almost unchanged the ancestry of their first Neolithic farmers. A shepherd leads sheep and goats, a woman grinds grain on a stone quern beside a potter, and a warrior with a horned helmet, a round shield and a bronze sword walks between round stone huts with thatched roofs. Clothing, buildings and the scene are approximations, and it is not a reconstruction of the individuals whose genomes are discussed here.

In 2008, John Novembre and colleagues showed that the genomes of 3,000 Europeans, summarised in two dimensions, draw a map of Europe: Iberians next to the French, Italians next to the Swiss, Poles next to the Germans. Genes mirror geography. But every genetic map of Europe also has a few populations that refuse to sit where geography puts them. Sardinians drift away from the rest of Italy. The Basques sit apart from their Spanish and French neighbours. The Saami of the far north, the Finns, the Roma and a handful of island peoples all pull away from the crowd.

Are these populations outliers for the same reason? Not at all. Some carry ancestry that the rest of Europe lost, others carry ancestry that the rest of Europe never had, and others simply drifted apart in isolation. We measured how distinctive each of 110 present-day populations of Europe is in Global25, and then tested what explains it.

The genetic map of Europe and its outliersPrincipal component analysis of 110 present-day populations of Europe (Global25 averages, our analysis)SardiniansBasquesCypriotsMalteseAshkenazi JewsRoma (Balkans)SaamiMariUdmurtsFinnsLatviansSorbsIberiaFranceBritish IslesScandinaviaBalkansRussia, UkraineGreece, AegeanVolga and UralsItaly, northItaly, southOff the chart: Kalmyks (East Asian ancestry)and Bashkirs (far below the Mari)PC1 (64% of variation): Mediterranean to northeastPC2 (22%): eastern and Asian ancestry (down)Outliers discussed hereItalian regionsOther European populations
Principal component analysis of present-day European population averages in Global25 (our analysis). Axes computed without Kalmyks, Roma, Ashkenazi Jews, Cypriots and Bashkirs, which are projected. Distances on this two-dimensional map are only a rough guide.

Measuring an outlier

A principal component plot is a good picture but a poor measuring tool: it keeps only two dimensions, and populations can look isolated on the plot without being so in the full genome. We used a stricter test. For each population, we tried to rebuild its Global25 profile as a mixture of all the other populations of Europe, excluding its own group (all Basques together, all Finns together, and so on). The leftover distance, the residual, measures what the rest of Europe cannot explain. Across the 110 populations the median residual is 5.9, and 90% of populations score below 18.

We then ran the same model a second time, adding outside and ancient sources: western hunter-gatherers, Neolithic Sardinians, Bronze Age and Iron Age Iberians, and present-day Siberian, South Asian, Levantine, North African, Han and Mongolian populations. If the residual collapses, the population is distinctive because it carries ancestry that the rest of Europe lacks. If it does not move, the population is distinctive because of drift: centuries of small numbers and isolation that reshuffle allele frequencies without adding anything new.

Before trusting the test, we checked it on artificial populations. When we added 5% of Siberian ancestry to the Tuscan profile, the residual rose from 3.0 to 7.9; at 10%, to 15.8; and the second model recovered 4.7% and 9.7% of Siberian ancestry. A residual therefore flags ancestry that is rare in the rest of Europe. It has one blind spot: when two populations share the same unusual ancestry, they partly hide each other. A Tuscan profile with 10% South Asian ancestry barely moved (4.6), because the Roma, who are in the pool, can absorb it.

How hard is each population to rebuild from other Europeans?Residual distance when modelled from all other European populations (red), then with outside and ancient sources added (green)020406080100What fixes itKalmyks250 (off scale)Mongol 92%Roma (Balkans)91South Asian 39%Sardinians73Sardinia Neolithic 69%Marinothing (drift)Saami41Siberian 11%Udmurtsnothing (drift)Basques (France)22Bronze Age Iberian-likeCypriots21Levant 47%Latviansnothing (drift)Finns (southwest)nothing (drift)Basques (Spain)19Bronze Age Iberian-likeFinns (east)nothing (drift)Sorbsnothing (drift)Maltese15Levant 21%, N. Africa 9%Icelanderslittle (drift)Ashkenazi JewsLevant 37%Italians (Lazio)Italians (Calabria)EnglishItalians (Tuscany)GermansOther Europeans onlyPlus outside and ancient sourcesShaded: typical Europe (median 5.9). Dashed line: 90% of populations score below 18. Purple: no outside source helps.
Residual distance (x 1000) when each population is modelled as a mixture of all other European populations, excluding its own group (red), and when outside and ancient sources are added (green). Right column: the source that explains the gap, or none (our analysis).

Twelve populations score above 18. They tell four kinds of story, plus one surprise.

Time capsules: Sardinians and Basques

The Sardinians are the most famous outliers of Europe, and our test shows why. Rebuilt from other Europeans, they leave a residual of 73, the third highest of the continent; their nearest neighbours, the Corsicans, sit 74 units away. Add Neolithic Sardinians as a source and the residual falls to 12, with 69% of the profile taken from people who lived on the island more than 5,000 years ago. Early and Middle Bronze Age Sardinians, before and during the Nuragic period, work just as well (74 to 77%), and so do the Early Neolithic farmers of Iberia (71%). Anatolian farmers do much worse (30), as do the Linear Pottery farmers of central Europe (24). The ancestry the Sardinians kept is specifically that of the western Mediterranean Neolithic.

The literature says the same thing. In 2018, Charleston Chiang and colleagues sequenced 3,514 Sardinians and found that they are the Europeans most similar to Neolithic farmers, with the strongest isolation in the mountainous Ogliastra, and some shared ancestry with the Basques. In 2020, Joseph Marcus and colleagues published 70 ancient Sardinians and found long genetic continuity from the Neolithic to the end of the Nuragic period, in the second millennium BC. The steppe migrations that transformed the mainland around 2500 BC barely reached the island. Change came later, from Phoenician and Punic times onward, with admixture from the eastern and northern Mediterranean, unevenly spread across the island.

The Basques are a younger time capsule. In 2019, Inigo Olalde and colleagues showed that steppe ancestry replaced about 40% of the Iberian gene pool by 2000 BC, and that from Roman times onward Iberia received gene flow from North Africa and the eastern Mediterranean. The Basques were the exception: in their words, a typical Iron Age population without the later admixture that affected the rest of Iberia. In our test, the French and Spanish Basques score 21.7 and 19.4. The Spanish Basque residual falls to 11 when Bronze Age Iberians are added, at 47% of the profile. In the seven-source model below, Basques carry 15% hunter-gatherer, 57% farmer and 28% steppe ancestry, and no North African component at all, where Castilians carry about 7%.

The Basques are not a complete island, though. Their neighbours are partly modelled from them: Navarra at 55%, La Rioja at 49%, Burgos at 31%. And the one surprise of our ranking, Asturias (21.3), is modelled with almost a quarter of Basque ancestry. Our reading, which a larger Asturian sample should test, is that the Basques are the most extreme point of a northern Iberian profile that the rest of Spain diluted more.

Each outlier next to its neighbourSeven-source model: hunter-gatherer, Anatolian farmer, steppe, plus four outside sources (our analysis)0%25%50%75%100%fitSardinians1082738Tuscans54341115Basques (Spain)15572845Castilians105231732Saami1510502552Swedes14335342Finns (east)17205659Estonians18255771Mari145029117Russians (Kostroma)13265661Roma (Balkans)2515183918Bulgarians514329Ashkenazi Jews30214314Poles14345257Maltese372232911Sicilians (west)39213015Cypriots1787516Greeks (Thessaly)53361117Hunter-gath.Early farmerSteppeSiberianLevantSouth AsianN. AfricanLevant = modern Lebanese, a stand-in for eastern Mediterranean sources. Fit: lower is better; northeastern fits are rough.
Each outlier paired with a neighbour, modelled with seven sources: Loschbour hunter-gatherer, Barcin Anatolian farmer, Samara Yamnaya, Nganasan, Lebanese, Gujarati and Moroccan (our analysis). Values in percent; fit distance x 1000 on the right.

Newcomers: Kalmyks, Roma and Ashkenazi Jews

The highest residual in Europe belongs to the Kalmyks of the lower Volga, at 250: no combination of European populations comes close. Add a Mongolian source and it drops to 12, with 92% of Mongol ancestry. The Kalmyks are Oirat Mongols who reached the steppe west of the Caspian in the early 17th century. In 2024, Natalia Balinova and colleagues showed that the Kalmyks and the Oirats of western Mongolia remain strikingly similar genetically, four centuries and thousands of kilometres apart.

The Roma of the Balkans come second, at 91, and their residual collapses to 11.5 once a South Asian source is available. The seven-source model gives them 39% South Asian ancestry, the rest being largely Balkan and West Asian. This matches the study of Isabel Mendizabal and colleagues in 2012: the Roma descend from a group that left north or northwest India about 1,500 years ago and spread across Europe from the Balkans about 900 years ago, mixing with local populations along the way.

The Ashkenazi Jews are the instructive case. Their residual is only 8.0, close to the European median, because Sicilians, Maltese and Cypriots in the pool can mimic their average profile. Yet Ashkenazi Jews are one of the most distinctive groups of Europe in medical genetics. In 2014, Shai Carmi and colleagues sequenced 128 Ashkenazi genomes, found an even mixture of European and Middle Eastern ancestry, and traced all Ashkenazi Jews to a bottleneck of about 350 people in late medieval times. That kind of distinctiveness lives in long shared DNA segments, which a summary like Global25 does not record. Our test sees ancestry; it does not see founder effects well.

The Siberian edge: Saami, Mari and Udmurts

At the northeastern end of the map, the Saami score 41, the Mari 47 and the Udmurts 27. For the Saami, an outside source helps: adding Siberian Nganasans halves the residual, to 20, and the seven-source model gives them 25% Siberian ancestry and only 10% farmer ancestry, the lowest of any European population except the Kalmyks. In 2018, Thiseas Lamnidis and colleagues found this Siberian ancestry already present on the Kola Peninsula around 1600 BC, at Bolshoy Oleni Ostrov, and showed that the Saami carry the most of it among the peoples of Fennoscandia. The same year, Kristiina Tambets and colleagues found that most Uralic speakers, from the Saami to the Volga, share a component of likely Siberian origin.

The Mari and Udmurts carry it too, 29% and 24% in our model, but adding a Siberian source barely changes their residual. Their distinctiveness is mostly drift, and partly a sampling problem: the populations closest to them, in the Volga and Ural region, are few in our pool. They sit where the genetic map of Europe meets that of Siberia.

Drift islands: Finns, Latvians and Sorbs

The Finns of the southwest and the east score 18.9 and 17.8, and adding outside sources changes nothing: 18.9 and 17.4. Their Siberian share is small, 4 to 7%. What sets them apart is history, not exotic ancestry. Finland was settled by a small number of founders and stayed isolated for centuries, and the late settlement of the east added further bottlenecks. In 2017, Sini Kerminen and colleagues found strong regional structure in Finland, with a main genetic border between west and east that follows the 1323 border of the Treaty of Noteborg between Sweden and Novgorod, which also separated the early and late settlement areas.

The Latvians (20.6) and the Sorbs (16.7) belong to the same family. Outside sources do not help them either. The Latvians carry the highest hunter-gatherer share of our sample, 19%, an old Baltic signature that their Estonian and Lithuanian neighbours share in slightly smaller doses. The Sorbs, a Slavic-speaking minority of eastern Germany, are closest to the Poles, which fits the study of Krishna Veeramah and colleagues in 2011: they look Slavic rather than German, and only a minor genetic isolate. Global25 tends to understate drift: the Icelanders, a classic founder population, score only 8.9.

Mediterranean crossroads: Cypriots and Maltese

The Cypriots (21.4) and, to a lesser degree, the Maltese (14.9) are edge populations too, on the southeastern side: islands close to other worlds. Adding a Levantine source brings the Cypriot residual down to 9.3, with 47% Levantine ancestry. The Maltese residual halves to 7.3 with 21% Levantine and 9% North African ancestry, the highest North African share of our sample, just ahead of western and central Iberia (7 to 8%). The Levantine proxy here is modern Lebanese, a stand-in for several eastern Mediterranean sources, ancient and modern, so these percentages measure a direction rather than a precise origin.

And the Italians?

Italy is often listed among Europe's genetic outliers, and with reason: in 2019, Alessandro Raveane and colleagues found in Italy the largest degree of genetic structure detected so far in the continent, with northern Italians close to other Europeans and southern Italians shifted towards the eastern Mediterranean. On our map, the distance from Lombardy to Calabria is almost as large as from Paris to Poland. But structure is not isolation. Every Italian region except Sardinia is easy to rebuild from its neighbours: Tuscany scores 3.0, Apulia 3.3, Campania 2.6, Calabria 5.5, eastern Sicily 9.0. Only Liguria (14.5) stands out a little, possibly a quirk of the sample.

Italy: a gradient, not an outlierAncestry by region from north to south (seven-source model), and how hard each region is to rebuild from other Europeans0%25%50%75%100%residualLombardy62368Piedmont58384Veneto58374Emilia57364Tuscany5434113Marche4830203Lazio4827238Abruzzo4427293Campania3922383Apulia4325323Calabria3821426Sicily (east)4222369Sicily (west)3921369Sardinia1082773Hunter-gathererEarly farmerSteppeEastern Mediterranean and otherRight column: residual from other Europeans only (median Europe 5.9). Only Sardinia stands out.
Italian regions from north to south, with Sardinia for comparison, in the same seven-source model; the four outside sources are grouped in red (our analysis). Right column: residual when each region is rebuilt from other Europeans.

What the Italian regions show is a smooth gradient. From north to south, the share modelled as eastern Mediterranean rises from almost nothing in Lombardy to about 40% in Calabria, while steppe ancestry falls from 36 to 21%. The exact numbers depend on the proxy, which probably overstates the eastern share, but the gradient does not. Ancient DNA dates part of it: in 2019, Margaret Antonio and colleagues found that the population of Imperial Rome shifted strongly towards the eastern Mediterranean. Italy is not a genetic island; it is a bridge between Europe and the eastern Mediterranean, with Sardinia as the one true exception.

Four ways to become an outlier

Put together, Europe's outliers tell four different stories. Time capsules kept an old ancestry that the rest of the continent diluted: the western Mediterranean farmers in Sardinia, the Iron Age Iberians in the Basque country. Newcomers brought an ancestry Europe did not have: the Kalmyks from Mongolia, the Roma from India, the Ashkenazi Jews partly from the Middle East. Edge populations sit where Europe meets Siberia or the Levant: the Saami, Mari and Udmurts on one side, Cypriots and Maltese on the other. And drift islands, like the Finns, Latvians and Sorbs, became distinctive simply by being few and isolated for a long time.

How Europe's outliers were madeTen events behind the populations that stand apart6000-5500 BCFarmers reach Sardinia and IberiaWestern Mediterranean Neolithic seafarers found the gene poolthat Sardinians still largely carry.2500-2000 BCSteppe ancestry sweeps EuropeAbout 40% of Iberian ancestry is replaced by 2000 BC;Sardinia is barely touched.c. 1600 BCSiberian ancestry reaches the Kola PeninsulaFound at Bolshoy Oleni Ostrov, near the Barents Sea:the ancestry the Saami carry most today.1st mill. BCPhoenicians and CarthaginiansEastern Mediterranean admixture begins on Sardinia's coasts.Roman eraThe Mediterranean mixesEastern Mediterranean and later North African ancestry reachItaly and Iberia. The Basque country is largely spared.c. 500 ADAncestors of the Roma leave IndiaA founding group from north or northwest India,about 1,500 years ago.c. 1100 ADThe Roma spread from the BalkansDispersal across Europe begins about 900 years ago.1200-1400Ashkenazi bottleneckAll Ashkenazi Jews descend from a founding groupof roughly 350 people, 600 to 800 years ago.1323Treaty of Noteborg splits FinlandThe border still matches the genetic divide betweenearly-settled west and late-settled east Finland.1630Kalmyks reach the VolgaOirat Mongols settle the steppe west of the Caspian,Europe's only Buddhist Mongol people.Sources: Marcus 2020, Olalde 2019, Lamnidis 2018, Mendizabal 2012, Carmi 2014, Kerminen 2017.
Key events behind the populations that stand apart on the genetic map of Europe. Dates are approximate.

Some limits should be stated. Global25 summarises the genome in 25 dimensions and loses most of the information carried by shared segments, so it detects ancestry better than drift and founder effects. Population averages hide variation inside each group, and some samples are small. Our outside sources are proxies, especially the modern Lebanese for the eastern Mediterranean and the modern Nganasans for Siberian ancestry. Finally, an outlier score is always relative to the pool: two populations that share an unusual ancestry partly hide each other, as the Basques and Sardinians, or the Sicilians and Ashkenazi Jews, do here.

Key points

  • Rebuilt from all other European populations, the median population of Europe leaves a residual of 5.9; twelve of 110 score above 18.
  • Sardinians (73) are a time capsule of western Mediterranean Neolithic farmers: Neolithic or Nuragic Sardinians explain about 70 to 77% of their profile.
  • Basques (19 to 22) are Iron Age Iberians who missed the later North African and eastern Mediterranean inputs, but their neighbours share part of their profile.
  • Kalmyks (250) and Roma (91) are outliers because they carry Mongolian and South Asian ancestry; Ashkenazi Jews are distinctive through founder effects that Global25 barely sees.
  • Saami, Mari and Udmurts carry 24 to 29% Siberian-related ancestry; Finns, Latvians and Sorbs stand apart through drift, not foreign ancestry.
  • Italians are not outliers: every mainland region and Sicily scores between 2.6 and 14.5, along a smooth north to south gradient towards the eastern Mediterranean. Only Sardinia stands apart.

Global25 coordinates used in this article

The outlier populations, their neighbours, the Italian regions and the outside and ancient sources. They can be pasted directly into the Vahaduo tool.

Sardinian,0.121687,0.167285,0.028490,-0.050652,0.060151,-0.022134,-0.003952,0.002496,0.041574,0.077351,-0.000059,0.016649,-0.028664,-0.012974,-0.013572,-0.003001,0.011403,-0.001347,0.001680,-0.012995,-0.002121,-0.001102,-0.010084,-0.021383,0.000337
Corsican,0.120327,0.151822,0.019530,-0.023002,0.035039,-0.010040,-0.001846,-0.000561,0.015705,0.033792,-0.000603,0.007461,-0.013231,-0.007107,-0.004537,-0.000597,0.004042,0.001249,0.002451,-0.003046,-0.000392,0.000247,-0.002579,-0.001386,-0.000530
Basque_Spanish,0.126424,0.148739,0.055679,0.008790,0.055450,0.000249,-0.001754,0.000025,0.029539,0.042923,-0.005214,0.010801,-0.024986,-0.019493,0.015807,0.002614,-0.005630,0.003172,-0.002375,-0.001791,0.008819,0.002204,-0.006479,-0.008654,0.001150
Basque_French,0.128051,0.152025,0.055173,0.012823,0.056411,0.002510,-0.001410,0.003323,0.030597,0.041204,-0.009272,0.010341,-0.020976,-0.014024,0.013219,-0.001936,-0.011917,0.003028,-0.000716,-0.004415,0.010157,0.002485,-0.008640,-0.009170,0.000072
Spanish_Navarra,0.119514,0.146236,0.044249,-0.000754,0.048727,-0.001208,-0.003212,-0.001461,0.024952,0.038695,-0.004980,0.011490,-0.015263,-0.017019,0.015653,0.002829,-0.008736,0.001098,-0.002472,-0.002543,0.004658,0.002556,-0.006943,0.000040,0.004750
Spanish_Castilla_Y_Leon,0.107753,0.143867,0.040477,-0.002153,0.043803,0.001859,-0.001723,0.004615,0.028429,0.029218,-0.002328,0.007843,-0.012091,-0.012019,0.012260,-0.006541,-0.006780,-0.001731,-0.004525,-0.002293,0.000291,-0.000907,0.001315,-0.002571,-0.002754
Spanish_Asturias,0.110408,0.144205,0.045632,0.000646,0.051702,-0.007251,-0.007285,0.006923,0.025975,0.031345,-0.003248,0.008093,-0.019029,-0.009083,0.014386,-0.013392,-0.015907,0.001267,-0.005405,-0.000875,0.006863,-0.006059,-0.001602,0.001325,0.006466
Saami,0.111262,-0.030212,0.111722,0.079781,-0.009329,0.008297,0.008137,0.013860,0.002966,-0.032096,0.022288,-0.007475,0.018071,-0.019826,-0.004004,0.002693,0.001157,-0.001489,-0.005986,0.004494,0.015816,0.001345,-0.002565,0.002658,0.000344
Swedish,0.132193,0.125885,0.072642,0.058972,0.040264,0.020790,0.006380,0.009294,0.003649,-0.006663,-0.004203,0.000533,-0.003045,-0.003352,0.014593,0.005865,-0.005106,0.001622,0.002991,0.005015,0.006773,0.002526,0.001253,0.011594,0.000212
Finnish_East,0.125855,0.078776,0.094388,0.083842,0.027390,0.020877,0.007554,0.013945,0.004587,-0.030590,0.007377,-0.007900,0.016225,0.002084,-0.004072,0.008902,0.011008,-0.001647,-0.001365,0.006021,0.009751,0.001131,0.001197,0.008005,0.002070
Finnish_Southwest,0.131323,0.096856,0.086313,0.072110,0.033737,0.022974,0.008549,0.014047,0.004474,-0.020889,0.003897,-0.005807,0.011465,0.008584,0.005293,-0.008569,-0.020112,0.001441,0.001587,-0.000219,0.006863,0.000526,0.002773,0.005121,0.003907
Estonian,0.132945,0.114146,0.087756,0.083302,0.042777,0.029758,0.011375,0.014446,0.000409,-0.028447,0.000065,-0.012349,0.019460,0.021359,-0.006664,0.001710,0.000222,-0.002483,0.001345,0.002214,0.001385,-0.001867,0.004264,0.000747,0.003054
Latvian,0.135449,0.122473,0.088774,0.089471,0.043208,0.034025,0.012597,0.014076,-0.002168,-0.036885,-0.003573,-0.013818,0.023221,0.031378,-0.010559,0.004057,0.002712,-0.000380,0.001609,0.004302,-0.002795,-0.005713,0.009071,-0.009327,0.002634
Sorb_Niederlausitz,0.130897,0.137096,0.068636,0.056848,0.050779,0.021196,0.011281,0.011076,-0.003477,-0.017312,-0.002111,-0.006744,0.013082,0.017891,-0.008415,0.001061,0.008475,-0.006208,0.007039,-0.004127,-0.003369,-0.007543,0.008011,-0.004458,0.001197
Polish,0.131840,0.129270,0.069868,0.057738,0.040676,0.021713,0.008678,0.010863,-0.000933,-0.018553,-0.004349,-0.006419,0.013093,0.018629,-0.007024,-0.000560,0.001546,-0.000074,0.002695,0.001287,-0.003129,-0.003119,0.005606,-0.003333,-0.000088
Icelandic,0.132698,0.133626,0.067819,0.056310,0.039366,0.022962,0.003349,0.008519,0.002642,0.000061,-0.005481,0.005170,-0.006368,-0.009083,0.023072,0.005293,-0.011691,0.001911,0.001739,0.003887,0.009255,0.002803,-0.000842,0.014018,0.000828
Mari,0.102441,-0.047730,0.093442,0.063057,-0.028723,0.001177,0.012142,0.016948,-0.004636,-0.039545,0.026235,-0.013621,0.032408,-0.038794,-0.035302,-0.023218,-0.001014,-0.009431,-0.039748,-0.032877,0.014266,0.004685,-0.048286,0.009881,-0.001504
Udmurt,0.107841,-0.020702,0.079718,0.066315,-0.030570,0.008875,0.008617,0.010716,-0.011055,-0.035615,0.021186,-0.005995,0.015072,-0.030880,-0.010391,-0.005453,-0.004182,-0.001000,-0.009682,-0.009588,0.000736,0.003152,-0.004352,0.008175,-0.002803
Russian_Kostroma,0.128241,0.087336,0.081458,0.068046,0.027441,0.021521,0.011907,0.012999,-0.003375,-0.029947,0.004222,-0.007493,0.017963,0.012340,-0.009138,-0.003911,0.000913,-0.001626,-0.001760,0.000479,0.000062,-0.001443,0.003595,-0.000602,-0.002056
Kalmyk,0.048261,-0.333500,0.061244,-0.023321,-0.047024,-0.026606,0.016075,0.015830,0.000777,0.008529,-0.031958,-0.003387,-0.002081,0.002064,0.002389,0.001830,-0.002921,0.002584,0.006159,0.011956,-0.018118,-0.008037,-0.014716,-0.001446,0.001581
Roma_Balkans,0.091058,0.063640,-0.058831,0.024979,-0.018773,0.013108,0.000392,0.002538,0.016703,0.014883,0.000379,-0.000350,0.000941,0.008762,-0.010541,-0.005215,0.008866,0.003125,0.001508,0.001000,-0.002080,0.000124,-0.000657,-0.004017,-0.001277
Bulgarian,0.124750,0.134862,0.025644,0.005362,0.024620,-0.000837,0.005499,0.003046,-0.000286,0.006998,-0.003345,0.000899,0.002111,0.008643,-0.014305,-0.002228,0.007093,0.002002,0.003142,-0.003702,-0.004617,-0.000544,0.002810,-0.002482,-0.001078
Ashkenazi_Poland,0.100117,0.136843,-0.007338,-0.043000,0.009797,-0.015409,-0.000881,-0.000894,0.008198,0.014275,0.001164,-0.000593,0.002880,0.000166,-0.004145,-0.001674,-0.003765,0.000032,-0.000807,-0.004064,-0.002678,-0.002272,0.001689,0.002169,0.001527
Maltese,0.091912,0.146490,-0.002498,-0.043403,0.013272,-0.017814,-0.004406,-0.000923,0.011095,0.020524,0.006150,0.000974,-0.002899,-0.000723,-0.004258,0.000878,-0.000407,-0.003769,-0.000487,-0.002361,-0.006145,-0.002597,0.002819,0.001190,0.002904
Cypriot,0.101086,0.146623,-0.035054,-0.068584,-0.003283,-0.023188,-0.000269,-0.003967,-0.001724,0.016792,0.003209,0.001313,0.000566,0.002497,-0.012506,0.000284,0.005110,0.000555,0.003220,-0.002257,-0.002151,0.001195,-0.001544,0.002668,-0.000998
Greek_Thessaly,0.119135,0.145221,0.013702,-0.020026,0.018875,-0.005113,0.000548,0.000769,0.000000,0.018770,0.002652,0.004096,-0.007284,0.005688,-0.009093,-0.004199,0.002130,-0.000549,0.004148,-0.005336,-0.006988,0.001278,0.003574,0.001406,-0.003353
Italian_Lombardy,0.122929,0.147886,0.026728,-0.015746,0.033468,-0.007460,0.002144,0.002135,0.009408,0.030479,-0.002497,0.007493,-0.015498,-0.005453,-0.002155,-0.005386,-0.004140,0.001140,0.002891,-0.004174,-0.003603,0.001020,0.000246,0.002937,-0.001961
Italian_Tuscany,0.118810,0.147784,0.013882,-0.021011,0.025294,-0.008938,-0.001063,-0.001626,0.005649,0.023239,-0.000309,0.006066,-0.011475,-0.003703,-0.002243,-0.000814,0.002154,0.000290,0.003334,-0.003829,-0.001967,0.001407,-0.000734,0.003196,-0.001226
Italian_Lazio,0.113140,0.152736,0.006788,-0.034303,0.021173,-0.010933,0.003243,-0.000277,0.006422,0.024237,0.000520,0.002548,-0.009574,-0.005230,-0.006732,0.002943,0.001043,0.000456,0.002690,-0.004227,-0.001922,0.004229,0.000567,0.000723,-0.000575
Italian_Campania,0.108263,0.148384,-0.008529,-0.043456,0.014038,-0.016154,-0.000777,-0.002201,0.004507,0.022100,0.000968,0.002265,-0.004980,-0.001440,-0.006259,-0.000806,0.002332,0.000000,0.002814,-0.003728,-0.003964,0.001907,0.000256,0.002104,0.000189
Italian_Calabria,0.103437,0.148140,-0.010324,-0.046997,0.012618,-0.015025,-0.002438,-0.001586,0.004448,0.021231,0.002213,0.002997,-0.004887,-0.001737,-0.006549,-0.002784,0.005802,0.000728,0.002938,-0.005362,-0.002605,-0.000325,0.000339,0.000015,-0.002425
Sicilian_East,0.103200,0.146574,-0.007291,-0.041129,0.017849,-0.015339,-0.001958,0.000077,0.007567,0.021322,0.007307,0.003897,-0.006145,0.003119,-0.005926,-0.006497,-0.002868,-0.001140,0.004483,-0.003877,-0.000042,0.002679,-0.001890,0.002289,0.000399
Sicilian_West,0.100164,0.145559,-0.000377,-0.035530,0.019388,-0.009854,-0.000627,0.000538,0.013362,0.022293,0.003410,0.003097,-0.002230,0.001881,-0.002262,-0.003182,-0.001869,-0.000042,-0.000880,-0.006128,0.000832,0.000824,-0.003204,-0.003494,0.000239
English,0.131855,0.137043,0.061788,0.044013,0.039230,0.016748,0.004985,0.005745,0.005264,0.005678,-0.004769,0.005616,-0.012589,-0.010351,0.020615,0.003552,-0.010369,0.004094,0.003685,0.002962,0.005937,0.003404,-0.003269,0.013794,0.000031
German,0.130299,0.137378,0.057363,0.037865,0.039566,0.015146,0.004177,0.005897,0.003746,0.001633,-0.004331,0.002316,-0.005334,-0.001744,0.008738,0.002822,-0.003300,0.001633,0.003531,0.001114,0.002947,0.001801,0.000337,0.008360,0.000074
Sardinia_Neolithic,0.124592,0.177015,0.043079,-0.058960,0.078997,-0.031064,-0.005369,-0.003018,0.062474,0.099613,0.003685,0.018341,-0.035267,-0.012979,-0.019982,-0.006007,0.008876,0.001930,0.005676,-0.011563,0.001852,0.002778,-0.012211,-0.029670,-0.000286
Sardinia_EBA_avg,0.125010,0.177026,0.045197,-0.051842,0.080431,-0.028087,-0.005004,0.001833,0.058612,0.095655,-0.002264,0.016041,-0.032383,-0.013758,-0.014703,-0.003681,0.008577,0.001523,0.003329,-0.008206,0.003766,0.003178,-0.012930,-0.029005,0.001531
Iberia_EBA_avg23,0.126146,0.154184,0.058896,0.000197,0.067772,-0.005105,-0.003873,0.002207,0.039455,0.056564,-0.003749,0.011507,-0.022920,-0.016604,0.003806,0.006462,0.008208,0.000408,0.000487,0.000114,0.009011,0.004978,-0.010438,-0.020144,0.002359
Iberia_IA_avg12,0.125680,0.147929,0.058045,0.004576,0.059293,-0.003161,-0.004602,0.004115,0.035706,0.045589,-0.002152,0.011202,-0.023129,-0.014691,0.010552,0.006254,0.006574,-0.001679,0.004169,-0.000552,0.005469,-0.000773,-0.004365,-0.012331,-0.000389
Loschbour_WHG,0.130897,0.109677,0.203645,0.198000,0.162492,0.059125,0.015041,0.038075,0.100217,0.016219,-0.015427,-0.017235,0.019921,-0.001239,0.061346,0.070670,0.002608,0.007348,-0.008925,0.065406,0.117543,0.010387,-0.049422,-0.173639,0.019519
Barcin_Anatolian_farmer,0.112685,0.182795,0.010936,-0.100130,0.055087,-0.046854,-0.002350,-0.002769,0.046836,0.081642,0.008607,0.010191,-0.015758,0.006193,-0.040173,-0.020021,0.003520,0.000507,0.011816,-0.014132,-0.008735,0.006306,-0.010969,-0.004940,-0.004670
Yamnaya_Samara,0.125838,0.089254,0.042908,0.115456,-0.027868,0.044685,0.004491,-0.002949,-0.054858,-0.072996,0.001858,0.000350,-0.001652,-0.023610,0.037263,0.015734,0.000000,-0.001478,-0.001704,0.012506,-0.003120,0.001374,0.011229,0.018436,-0.004524
Nganasan,0.044992,-0.411402,0.155635,0.000350,-0.160996,-0.090190,0.028671,0.041300,0.029406,0.013354,0.101939,0.009100,-0.004018,-0.026580,-0.020354,-0.010464,0.001934,0.013971,0.025496,-0.000886,0.042481,-0.012956,0.032291,0.000321,0.013000
Gujarati,0.051391,-0.057733,-0.156053,0.110822,-0.080584,0.062193,-0.000223,0.011607,0.033000,0.018852,-0.006496,0.000082,-0.002178,0.001741,-0.000679,0.000378,0.000880,-0.000171,-0.001427,-0.003933,0.001429,-0.003314,0.001109,0.001127,-0.002976
Lebanese_Christian,0.088149,0.145785,-0.048858,-0.078848,-0.012891,-0.030740,-0.000496,-0.004282,0.005181,0.010225,0.006658,-0.004529,0.010390,0.000000,-0.010315,0.008471,-0.002101,0.001788,0.002542,-0.000125,0.000763,-0.000302,-0.001410,0.002356,-0.001211
Moroccan,-0.085367,0.130213,-0.012822,-0.070881,0.021132,-0.032940,-0.027783,0.009769,0.055403,0.024399,0.009509,-0.009025,0.026990,-0.014466,0.016302,-0.006188,-0.002767,-0.017033,-0.038422,0.009435,-0.009844,-0.029512,0.020911,-0.001125,0.005921
Mongol,0.042088,-0.360259,0.060708,-0.030516,-0.042903,-0.024384,0.014961,0.016609,0.000051,0.007335,-0.033902,-0.003464,0.000419,0.001664,0.001598,-0.000569,-0.003126,0.000225,0.004191,0.009945,-0.012623,-0.006587,-0.013092,-0.001630,0.001432

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