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Ukrainians: Seven Thousand Years at the Crossroads of Steppe, Forest and Sea

§ 01

In 513 BCE, Darius the Great brought seven hundred thousand men into the Ukrainian steppe — and left with nothing. In 1971, a Ukrainian archaeologist found a Scythian gold pectoral ten centimetres from a looted burial chamber. The Trypillia people built the largest settlements in prehistoric Europe and systematically burned them every eighty years. The genetics of these lands is as layered as their history.

In 513 BCE, the Persian king Darius the Great — ruler of the largest empire the world had seen to that point, stretching from Egypt to the Indus — crossed the Bosphorus with his army and marched it into the steppes north of the Black Sea. By Herodotus's account, seven hundred thousand men marched under his command. The political objective was clear: to punish the Scythians for their earlier incursions into Persian territory and bring them under imperial control.

The Scythians offered no engagement. They retreated, burning their own fields, filling their wells, driving off their herds. The Persian army advanced towards the Don — and found no one to fight, nothing to capture. Darius sent the Scythian king Idanthyrsus a message: either fight, or submit to his sovereignty. Idanthyrsus replied that the Scythians had no cities to defend and no crops to protect, which meant that it was not he but Darius himself who would determine when and whether a battle occurred. Eventually Darius turned back, having won no engagement, taken no territory, and lost a substantial portion of his army to exhaustion and hunger.

Herodotus recorded this campaign in the fourth book of his Histories, having personally visited the Greek city of Olbia on the Bug estuary — on the territory of modern Ukraine — to gather firsthand accounts. His notes constitute the earliest detailed description of the people who inhabited this region. Genetics examines the same people through an entirely different lens — and the picture it reconstructs is considerably more complex than any literary account of a 'people'.

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The deepest layer: the hunters of the Dnieper

The genetic history of Ukrainian territory begins at the Dnieper Rapids — where the river cuts through outcrops of ancient crystalline rock and forms cataracts that rendered it impassable to boats for millennia. It is here that some of the earliest well-preserved genomes from the region have been found. A 2017 study in Current Biology analysed two men from the Dnieper Rapids area — one Mesolithic, one Neolithic — separated by roughly four thousand years. In the coordinate system of population genetics, they formed a remarkably close pair: despite a temporal gap larger than the one separating us from ancient Egypt, their genomes were genetically very similar. Everything around them was changing — agriculture was advancing from the west, new groups arriving from the east — yet along the rapids, the genetic line persisted.

The same Nature (2018) study adds a detail that illustrates the complexity well: a single Neolithic individual from Deriivka, dated to approximately 4950–4800 BCE, carries a fully Anatolian Neolithic genetic profile — genetically indistinguishable from a 'classic' Near Eastern farmer, despite living thousands of kilometres from Anatolia. Alongside him were people with entirely different origins. The 'Neolithic' on Ukrainian territory was not a single demographic scenario but a mosaic of groups existing and interacting in parallel.

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The Trypillia culture: the largest settlements in prehistoric Europe

From approximately 5500 to 2750 BCE, a culture known today as Trypillia or Cucuteni–Trypillia existed across the territory of modern Ukraine, Moldova and Romania. It produced what researchers cautiously call 'mega-sites': settlements of up to 350 hectares, with populations estimated at between ten and fifteen thousand people. This is comparable to the largest cities of Mesopotamia in the same period — but without monumental temples, without writing, without any obvious centralised authority.

Houses were arranged in concentric rings, with streets leading to a central open space. And every sixty to eighty years, the settlements were systematically burned and rebuilt — the even distribution of ash layers suggests this was a deliberate process. Why it was done remains one of the open questions of European prehistory: ritual renewal, sanitary considerations, or a social mechanism for redistributing power. There is no answer. Genetically, Trypillia people had mixed origins — an Anatolian farming substrate combined with local hunter-gatherer lineages. The Nature (2025) study of the North Pontic region shows that it was precisely Trypillia groups who came into contact with the steppe populations from whose mixing the Yamnaya genetic profile would subsequently emerge. Around 3000–2750 BCE, Trypillia settlements disappeared across their vast range with striking synchrony.

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The North Pontic steppe as a source of demographic impetus for the whole of Europe

If there is a place on the European map where the genetic component that most radically altered the continent's demographic profile in the Bronze Age was being assembled, it is the steppes north of the Black Sea. Two major Nature studies (2023 and 2025) reconstruct this process in detail unavailable even a decade ago. Around 4500–3500 BCE, a process of mixing was under way in the Ukrainian steppe between late farmers and groups with genetics inclined towards Caucasian and Eastern European hunter-gatherer substrates. Through several intermediate stages — the Sredny Stog and Usatovo cultures — the 'Yamnaya profile' crystallised, and around 3300 BCE a large-scale expansion began westward and eastward. In Corded Ware samples — the direct successor of this Yamnaya expansion — the steppe component constituted 70–80 percent of ancestry. What happened in the Ukrainian steppes in the fourth millennium BCE altered the genomes of much of Europe within a few centuries.

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Scythians: an ethnonym as a political designation, not a genetic category

In 1971, Ukrainian archaeologist Borys Mozolevskyi was excavating the Tovsta Mohyla kurgan in Dnipropetrovsk Oblast. On 21 June, clearing a burial shaft, he felt something scratch his finger. Under a thin layer of clay lay a gold pectoral of a Scythian king from the fourth century BCE: a crescent 30.5 centimetres in diameter weighing over a kilogram, made by Greek goldsmiths in three registers of scenes — animals, herdsmen, mythological creatures. Ancient grave robbers had passed ten centimetres from it without finding it. Mozolevskyi transported it to Kyiv concealed under an old padded jacket. It is now in the Museum of Historical Treasures of Ukraine in Kyiv.

This object is not only a work of art. It was made by a Greek craftsman for a Scythian king, with motifs drawn from Scythian daily life. It already contains the complexity that genetics reveals: 'Scythian' society was an interweaving of people of different origins, united by political structure, way of life and commercial ties with the Greek world — but by no means by shared ancestry. Herodotus himself recognised this. 'Scythians' was the Greek designation for many different groups: the 'Royal Scythians' of Crimea who constituted the political elite; the 'agricultural Scythians' of the middle Dnieper who appear never to have been nomads; the 'nomadic Scythians' of the open steppe. It was a Greek label for a broad political-cultural zone, not a self-designation of a single unified people.

The most detailed genetic study of Ukrainian territory — Science Advances (2025), covering 91 ancient genomes from 7000 BCE to 1800 CE — confirms this picture. The genetics of the Iron Age is heterogeneous. Some samples from different regions of Ukraine stand close to the Eastern European genetic continuum and to modern Ukrainians. Others show pronounced Central Asian or Siberian affinity — particularly among samples associated with wealthy kurgan burials. In one pre-Scythian sample there appeared the Y-chromosome lineage Q1b-YP4004, more typical of Central Asia. The rich kurgan elite and the ordinary population often had different genetic profiles, united by political organisation rather than common descent.

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Cumans, the Golden Horde, Nogais: the eastern signal was real, but limited in scope

The same Science Advances (2025) study allows a precise assessment of how substantially the subsequent waves of steppe nomads — Sarmatians, Huns, Khazars, Cumans, the Mongol Golden Horde — altered the genetics of the Ukrainian population. Late medieval nomadic burials identified on Ukrainian territory do carry high proportions of East Asian ancestry — in some samples 40–60 percent. This is a real and measurable genetic trace of the Mongol Horde, Cumans and Nogais. However, genetics registers them as distinct migrant communities of the steppe zone, rather than as broad influence on the settled population. When samples from the same time periods identified as 'White Croats' and early Cossacks — groups associated with the settled population of the region — are analysed, they stand close to the clusters of modern Ukrainians, with minimal East Asian signal. The steppe political history of Ukraine and the genetic history of most of its inhabitants are not the same thing.

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The early Middle Ages: northern Ukraine in models of early Slavic origins

The largest study to date on the genetics of early Slavic expansion, published in Nature (2025), used spatial models to reconstruct the probable area of origin of the early Slavic genetic profile. According to its findings, the area that best explains the origin point of this profile is southern Belarus and northern Ukraine. From here, by the genetic evidence, a demographic impulse beginning in the fifth to seventh centuries CE transformed the genetic profile of much of Central, Eastern and Southern Europe over the course of a few centuries. For Poland and north-western Ukraine, the authors estimate the replacement of the local gene pool by the early Slavic component at 93 ± 3 percent — a figure that captures the scale of the early medieval demographic transformation.

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Modern Ukrainians: their position on the genetic map

Among Y-chromosome lineages in modern Ukrainians, R1a (around 43%) and I (around 27%) dominate — lineages with deep roots in Eastern Europe, present in both Bronze Age steppe samples and early medieval Eastern European populations. East Asian Y-lineages characteristic of certain Turkic and Mongolic peoples are minimally represented in the modern Ukrainian population. The mitochondrial profile is typically European, with haplogroup H and other Western European lineages dominating. By autosomal DNA, Ukrainians form a cluster with Poles, Belarusians and other Eastern and Central European populations. A regional gradient exists within the country — western, central and eastern regions differ somewhat in component proportions — but this is a gradient, not a boundary.

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Conclusion

Darius returned from the Scythian campaign with nothing — and this fact entered history as one of the earliest documented defeats of a 'great army' by a tactic that had no name, because it was not a military strategy in the conventional sense: simply moving to where the enemy cannot follow. Herodotus went to Olbia and recorded what he heard. Borys Mozolevskyi in 1971 found a pectoral ten centimetres from a looted chamber — and that object, made by a Greek for a Scythian, is itself a precise metaphor for what Ukrainian territory has been across millennia: a place where different cultures not merely collided but produced something new.

The genetic history of Ukrainian lands is seven thousand years of continuous movement, mixture and renewal. Hunters of the Dnieper Rapids, preserving their lineages through Neolithic waves. Trypillia builders with their systematically renewed mega-settlements, who became part of the demographic process that gave rise to the steppe expansion. Yamnaya pastoralists whose genetic signal spread across most of Europe within a few centuries. Genetically heterogeneous 'Scythians', united by political structure rather than shared origins. Medieval steppe nomads with pronounced East Asian signals — real, but demographically limited in their impact on the settled population. And the early medieval Eastern European population from northern Ukraine — the one that formed the genetic profile that persists in modern Ukrainians.

This is not the history of one people. It is the history of one of the most intensively inhabited crossroads of Eurasia — with a continuous thread running through every succession.

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Glossary

WHG / EHG (Western and Eastern European Hunter-Gatherers) — two genetically distinct clusters of pre-farming European populations. Ukrainian Mesolithic samples tend towards the 'eastern' end of this spectrum.

Anatolian Neolithic component — the genetic heritage of the first farmers who arrived from the Near East around 8–9 thousand years ago. Present in modern Europeans, including Ukrainians, in varying proportions.

Steppe component / Yamnaya profile — the genetic heritage of Eurasian steppe pastoralists who migrated into Europe around five thousand years ago. The North Pontic region is one of the key zones where this profile was assembled.

R1a — a Y-chromosome haplogroup dominant in modern Ukrainians (around 43%). Widely found in Bronze Age steppe samples and early medieval Eastern European populations.

qpAdm — a statistical method in population genetics for estimating the proportional contributions of several 'source' populations to a 'target' population. Used in studies of Yamnaya expansion and early Slavic migrations.

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