The Formation of Modern Phenotypes

Neolithic Europe (7000-5000 BC)

Western Hunter-Gatherers

Western Hunter-Gatherers (WHG), also called West European Hunter-Gatherers or the Oberkassel cluster, were a Mesolithic population (c. 15,000–5,000 BP) that spread across western, southern, and central Europe after the Last Glacial Maximum. They are closely associated with the Villabruna cluster, a genetic lineage originating likely in the Balkans and expanding into Italy and Iberia by ~19,000 BP. WHGs expanded across Western Europe around 14,000–12,000 BP, replacing the earlier Magdalenian populations descended from Cro-Magnon settlers.

WHGs were one of two major postglacial hunter-gatherer groups in Holocene Europe, alongside Eastern Hunter-Gatherers (EHG). The border between WHG and EHG groups ran from the lower Danube up to the western Baltic. In Scandinavia, Scandinavian Hunter-Gatherers (SHG) formed as a mix of WHG and EHG. In Iberia, early Holocene hunter-gatherers were a mix of WHG and Magdalenian ancestry (e.g., GoyetQ2).

WHGs were mostly replaced by Early European Farmers (EEFs) of Anatolian origin during the Neolithic, though some WHG ancestry persisted via admixture. Today, the highest WHG ancestry is found in eastern Baltic populations.

Phenotypically, WHGs likely had dark skin and light eyes, lacking the genes for the lighter skin tones typical of modern Europeans.

Chalcolithic Europe (5000-2000 BC)

The Yamnaya

The Yamnaya, also known as the Corded Mediterranid

Origins, Expansion, and Legacy

The Yamnaya culture, emerging around 3300 BC on the Pontic-Caspian steppe, represents one of the most transformative populations in Eurasian prehistory. Genetically, physically, and linguistically, the Yamnaya people were the result of a unique synthesis of two major Upper Paleolithic-derived groups: Eastern Hunter-Gatherers (EHG) and Caucasus Hunter-Gatherers (CHG). This admixture occurred around 5000 BC on the eastern steppe, forming the genetic base known in archaeogenetics as Western Steppe Herder (WSH) ancestry. Later, minor gene flow from Anatolian and Levantine farmers further enriched this gene pool, though Early European Farmer (EEF) ancestry proper was notably absent due to the lack of Western Hunter-Gatherer (WHG) input.

 

Phenotype and Physical Characteristics

Anthropologically, the Yamnaya were tall, robust, and dolichocephalic, with large cranial capacity and prominent facial features. They typically had intermediate to light brown skin, wavy brown to black hair, and deep-set brown eyes, though genetic traces of blond hair mutations (from the Ancient North Eurasian Afontova Gora group) were beginning to emerge in some individuals. They were heavily built (mesomorphs), often with broad chins, strong jawlines, leptorrhine (narrow and often aquiline) noses, and pronounced brow ridges. Their high body hair and thick features likely reflected their rugged steppe environment.

 

Y-DNA and Genetic Legacy

The dominant paternal lineage among the Yamnaya was R1b-Z2103, a subclade of R1b-L23, distinct from the R1b-P312 lineage later prevalent in Western Europe. Some Yamnaya males also carried I2 lineages, and mtDNA showed a diversity of haplogroups including U, T, and others associated with CHG and early farmers. Intriguingly, a few Yamnaya specimens carried East Asian-associated mtDNA (C4), revealing a broad steppe network even at this early stage.

Despite their enormous autosomal impact, the Yamnaya paternal lineages rarely persist in modern Western Europe, suggesting elite dominance and founder effects played a major role in shaping subsequent societies without necessarily establishing a direct male-line continuity. This disparity between autosomal and Y-DNA patterns was noted especially in the Corded Ware culture, which had up to 75% Yamnaya-like ancestry, but predominantly R1a-M417 males, not R1b-Z2103.

 

Cultural Expansion and Linguistic Impact

The Yamnaya were likely the main vector for Indo-European languages west of the Caucasus, particularly the core Indo-European branch. Their migration westward into the Carpathian Basin and Danube valley between 3100–2600 BC helped trigger the formation of the Corded Ware culture, a hybrid society shaped in the contact zone with Neolithic groups. This culture would spread across Central and Northern Europe, becoming the ancestor of various later Indo-European-speaking peoples, especially those associated with R1a lineages.

The Catacomb, Poltavka, Srubnaya, and Sintashta cultures followed in the Yamnaya’s wake, expanding their genetic and linguistic influence into Eastern Europe, Central Asia, and eventually South Asia, particularly among Indo-Aryan populations. This steppe ancestry, primarily derived from Steppe_EMBA (Early-Middle Bronze Age) and Steppe_MLBA (Middle-Late Bronze Age) sources, accounts for up to 30–50% of the DNA of some modern South Asian populations, especially in the northwest.

In contrast, Southern Europe shows only moderate Yamnaya-related ancestry (18–33%), and Sardinians and Sicilians retain far less, reflecting their deeper connection to earlier Neolithic farmer lineages.

 

Yamnaya and Proto-Indo-European Origins

The linguistic impact of the Yamnaya remains central to the debate over the origin of the Proto-Indo-European language (PIE). The prevailing Kurgan hypothesis (Gimbutas, Anthony, Reich) situates the PIE urheimat in the Pontic-Caspian steppe, while newer proposals (e.g. Lazaridis 2024, Kroonen et al. 2022) refine this to the Lower Volga-Caucasus (CLV) region, differentiating core Indo-European from Indo-Anatolian, the latter perhaps arising further south or in Sredny Stog.

The Yamnaya language, therefore, may have been a late PIE dialect descended from an EHG substrate enriched by CHG/Caucasian phonology and morphology. Genetic clustering supports the idea of a socially stratified Yamnaya elite, dominated by a small group of males who spread both their genes and language far beyond their homeland. Nevertheless, the divergence between paternal lineages in Yamnaya and their supposed descendants (e.g., Corded Ware) raises questions about exact mechanisms of cultural and linguistic transmission.

 

Legacy and Modern Impact

Though the Yamnaya “type” may be extinct in the pure form since 2600 BC, their biological legacy is profound, contributing significantly to the height, lactose tolerance, and phenotypic variation of modern Europeans, especially in the north and east. Genetically, they introduced Ancient North Eurasian ancestry into Europe, forming one of the key pillars of the modern European gene pool alongside Western Hunter-Gatherers and Early Farmers.

Phenotypically, descendants of the Yamnaya—through recombination with Neolithic and Mesolithic populations—gave rise to several recognizable types:

• Corded Nordid in the east and north, with more gracile features.
• Pontid, particularly in Southeastern Europe.
• Keltic Nordid, evolving from interactions with Neolithic and megalithic populations in Western Europe, particularly under the influence of later Bell Beaker migrations.

Post-Yamnaya Steppe Morphology

Initially, populations of the post-Mariupol’, Sredniy Stog, and early Pit-Grave cultures in the Dnieper-Don-Donets region were characterized by massive, broad-faced proto-Europoid features. With the emergence of the Timber-Grave (Srubnaya) culture, the Ukrainian population represented a blend between the narrow-faced, dolichocephalic type associated with the Multi-Roller Ware (Babino) culture and the more massive broad-faced types of the Timber-Grave population from the Volga. Anthropological data suggest a clear influx from the Volga into Ukraine during the formation of the Timber-Grave culture, reinforcing the idea of population movements shaping regional types.

As the Timber-Grave culture progressed into the Belozerka stage, narrow-faced, dolichocephalic types became dominant, a trend that continued into the Scythian period. This morphological continuity indicates a genetic link between the Iranian-speaking Scythians and the earlier Timber-Grave population in Ukraine. The Pit-Grave culture, itself an heir of the Neolithic Dnieper-Donets and Sredniy Stog cultures, maintained a tall stature and robust Europoid cranial features.

Further east, the Abashevo culture, emerging in the forest-steppe zone alongside the Poltavka culture, was marked by narrow-faced dolichocephaly. The Abashevans had roots in the Balanovo and Fatyanovo cultures of the Middle Volga and also shared ancestry with Central European populations. The early Timber-Grave population, especially the Potapovka phase, was a result of admixture between a massive, broad-faced Pit-Grave–Poltavka type and a dolichocephalic Europoid type related to the Sintashta culture. Another distinct group participating in the steppe’s ethno-cultural dynamics was the Pokrovskiy type, which exhibited dolichocephalic, narrow-faced traits similar to the Abashevans but distinct from the Potapovkans.

Overall, the majority of Timber-Grave skulls are dolichocranic with medium-broad faces, pointing to the significant role of Pit-Grave and Poltavka ancestry. There is also evidence of a genetic connection between the Timber-Grave population of the Urals and the Alakul culture of the Urals and western Kazakhstan, both of which share morphological traits with the Sintashta people. Finally, the western segment of the Andronovo culture population also displays a dolichocranic cranial type, aligning it closely with the Timber-Grave culture, further underscoring the widespread diffusion and continuity of this narrow-faced Europoid lineage across the steppe.

The Corded Ware

Origins:

The Norid phenotype, often described as a Dinaricized Nordic type, is believed to have formed in Central Europe during the Bell Beaker period. It arose when R1b-bearing Indo-European steppe populations, carrying an admixed Yamnaya-derived morphology, migrated into Central Europe and intermingled with local Corded Ware populations. In the lower Rhine and surrounding regions, this mixture interacted with local Borreby elements, producing a tall, robust, and sharply featured type that moved into northern France, the Low Countries, and the British Isles. This population would eventually contribute to the formation of the Paleo Atlantid, Brünn, Keltic Nordid, North Atlantid, and Cymrid phenotypes, among others.

British Bell Beaker man

Upon migrating to the British Isles, this Bell Beaker population, known as the Bell Beaker Borreby, would overturn over 90% of the autosomal DNA and Y-Haplogroups of the initial EEF peoples. However, in the western regions of the two islands, the original population had a greater genetic influence on the newcomers. The result of this mixture, characterized by strong cranial features, wide faces, and deep-set eyes, is still visibly represented in parts of western Ireland, western Scotland, and Wales, where relative geographic isolation helped preserve this ancient substrate to the present. 

The Paleo Atlantid phenotype

A related branch of the Bell Beaker migratory movement pushed westward through central and southern France, where it absorbed more Early European Farmer (EEF) and Western Hunter-Gatherer (WHG) genetic input. Over time, in the isolated valleys and rugged terrain of the western Pyrenees, this lineage gave rise to the Baskid phenotype. Characterized by high foreheads, prominent noses, and strong facial bones, this type became relatively stable and endogamous in the Basque region, where ancient pre-Indo-European genetic and linguistic traits have persisted into the present.

Following the Bell Beaker phenomenon, a second wave of migrations from Central Europe introduced populations with more Dinariform traits, characterized by taller stature, narrower faces, and prominent nose bridges. This new element mixed with the existing Cro-Magnid populations in the British Isles and the Netherlands, resulting in various genetic and phenotypic clines. This admixture gave rise to the Keltic Nordid phenotype—a tall, mesocephalic type with harmonious features that became dominant in much of England and the Netherlands. Meanwhile, a more Cro-Magnid-influenced variety arose in Scotland and Eastern Ireland as more residual elements reached these regions. The more isolated western regions of Ireland retained the greatest degree of original Cro-Magnid morphology, known as the Paleo Atlantid phenotype.

Genetics: