European M406:
The Northern Route

That M406 was spread into Europe via the Mediterranean vectors of Greece and Rome is a fairly obvious conclusion. Especially with aDNA revealing Roman M306 samples with Levantine and Anatolian autosomal affinities. The genetic impact of such arrivals to Rome as slaves or low class immigrants is well documented and therefore wherever the Romans went in the days of the Empire, so to did G-M406.

For several years however, it has been clear that G-M406 also made its way into the continent via a northern route over the Caucasus mountains. As new aDNA has become available, it shows that men bearing the G-M406 signature were established on the Pontic Steppe. By around 9th century BC some were clearly assimilated into the nomadic Cimmerian and Scythian cultures who entered the region from Central Asia. Ultimately, these cultures and the other successive nomadic groups that would succeed them, served as conduits of G-M406 men into Eastern Europe and perhaps beyond.

From Anatolia to the Caucasus

It is well documented in aDNA and historical context that the epicenter of G-M406 in the Bronze Age was Central Anatolia. With many other ancient Haplogroup G subclades being associated with early Neolithic farming sites and found amongst early Neolithic sites in Europe, it remains clear that G-M406 were not among the First Farmers of Europe. As Ted Kendall is known to say, they "missed the boat," likely remaining hunter-gatherers in the forests of Central Anatolia for several millennia to come. Some eventually settled among in small villages associated with copper mining among the Pre-Hattic culture of Anatolia (see several G-M406 and G-M3302 samples in Genomic History of Neolithic to Bronze Age Anatolia (Skourtanioti et al., 2020). Many G-M406 men may have lived along Anatolia's Black Sea coast, later incorporated into the Hittite kingdom that by all accounts incorporated the native-Hattic peoples into its non-ruling classes. During the Middle and Late Bronze Age, G-M406 was also likely found among the rebel Kaska peoples and perhaps among the people of Lake Van who were the forerunners of the Armenians.

Evidence from both modern and ancient DNA show that as late as the Roman Empire, G-M406 men from Anatolia entered the Caucuses. With findings of Iron Age Cimmerian and Scythian G-M406 on the Pontic Steppe it is clear that the signature arrived much earlier in time. The Genetic History of the South Caucasus has previously concluded that Anatolian migrants contributed to the South Caucasus gene pool as early as the Middle Bronze Age, indicating that interactions between the regions were longstanding and likely facilitated by trade and geographic proximity. This suggests ample opportunity for G-M406 to have seeded itself amongst the Caucasus during the height of Bronze Age civilization, and also possibly as it began to collapse circa 1200 BC.

Assimilation on the Pontic Steppe

Some men bearing the G-M406 signature must have made their way out of the Caucuses and onto the Pontic Steppe by the time the Cimmerians arrived from Central Asia in 1000 BC.  The Cimmerians were the first of many Iranic horse nomads to arrive in the region. To some, the Cimmerians were simply the westernmost representatives of the Scythian culture. In fact, they seemed to have shared a common language, origins and culture with the Scythians.  

Still, archeologists often associate the Cimmerians with the pre-Scythian Chernogorovka-Novocherkassk complex. In addition, most sources from the time period clearly distinguished the Cimmerians and the Scythians as being two separate political entities. By all accounts the Cimmerians were displaced by the arrival of the Scythians, with some groups leaving the Steppe and crossing the Caucusus to raid and control part of Anatolia during the 7th century BC. 

Chernogorovka-Novocherkassk objects have also been found in the eastern parts of Central Europe, especially in Transylvania and on the Hungarian Plain. To some scholars, this indicates that the Cimmerians also made their way west along the Black Sea inspiring what some scholars call the Thraco-Cimmerian culture. Thraco-Cimmerian goods have been found spread further west over most of Eastern and Central Europe, as far as Denmark and eastern Prussia. 

According to researches such as Jan Olbrycht and Jan Bouzek, evidence of the Chernogorovka-Novocherkassk complex is  spread across a large area ranging from north-eastern Bulgaria and Moldova in the west through Ukraine and Crimea and up to the Kuban and Volga-Kama regions in the east. The Cimmerians therefore influenced an area ranging from Central Europe and the Hungarian Plain in the west to the Pontic and Ciscaucasian Steppes in the east.

Arrival of the Sythians

The Scythians were an ancient Eastern Iranic horse-riding nomadic people who migrated during the 9th to 8th centuries BC from Central Asia to the Pontic Steppe in modern-day Ukraine and Southern Russia. Skilled in mounted warfare, the Scythians became the dominant power on the western Eurasian Steppe in the 8th century BC. In the 7th century BC, the Scythians crossed the Caucasus Mountains and often raided West Asia along with the Cimmerians. In the 6th century BC, they were expelled from West Asia by the Medes, retreated back into the Pontic Steppe, and were later conquered by the Sarmatians in the 3rd to 2nd centuries BC. By the 3rd century AD, the last remnants of the Scythians were overwhelmed by the Goths, and by the early Middle Ages, the Scythians were assimilated and absorbed by the various successive populations who had moved into the Pontic Steppe.

The Sarmatians, Goths, and Huns

Following the Scythian era, the Sarmatians, an ancient Eastern Iranic nomadic confederation, emerged as the dominant power on the Pontic Steppe by the 3rd–2nd centuries BC. Descended from the Sauromatians and absorbing elements of the earlier Scythian population, the Sarmatians (including subgroups such as the Alans) controlled vast territories from the Don River in the east to the Danube in the west. Like their predecessors, they were skilled horse warriors whose culture emphasized mobility, warfare, and assimilation of conquered or allied groups. 

By the 1st century BC, the Sarmatians had pushed westward, clashing with and sometimes allying with the Roman Empire. Roman sources record large Sarmatian contingents serving as auxiliary cavalry; in AD 175, for example, Emperor Marcus Aurelius settled 5,500 Iazyges in Britain after the Marcomannic Wars. Archaeological and isotopic evidence confirms the presence of individuals with Pontic-Caspian and Caucasus-related ancestry in Roman Britain, demonstrating the extraordinary reach of Sarmatian mobility. Crucially for the G-M406 story, Sarmatian groups had already incorporated Caucasian and Anatolian-derived lineages during their earlier movements along the Pontic Steppe and through interactions with Alanic and other Iranic tribes. Ancient DNA confirms that men carrying the G-M406 signature (specifically the FGC5089 and M3302 branches already present among Scythians) were assimilated into Sarmatian society as a minority paternal lineage. These men were not part of any “core” Sarmatian genetic signal but traveled with them as warriors, allies, or assimilated subjects. The presence of a Roman-era Pannonian G-M406 sample shows how the Northern Route extended deep into Central Europe through Sarmatian networks.

The later arrival of the Goths (Germanic-speaking migrants who displaced or absorbed many Sarmatian groups in the 3rd–4th centuries AD) did not erase these lineages. Instead, the Goths served as another vector, carrying mixed Pontic-Steppe populations (including residual Sarmatian/Alanic elements) into the declining Western Roman Empire and the Carpathian Basin. By the time the Huns arrived in the late 4th century, the stage was already set for G-M406 to survive as a rare but persistent marker among the multi-ethnic steppe confederations that would follow.

The Avars and Magyars

The Avars (AD 580–804) replaced the Huns as the dominant force on the Pontic Steppe and along the Byzantine frontier. Like the Huns, their elite core originated in Central Asia with strong genetic ties to Xiongnu populations (as shown by Maróti et al. 2022). Their language was likely Turkic or Mongolic, and they arrived in the Carpathian Basin around AD 567 as a multi-ethnic confederacy. 

The majority of the Avar-period population, however, consisted of local European residents or earlier steppe remnants (Sarmatians, Alans, and others) absorbed during their westward migration.  These incorporated groups brought branches of G-M406 into the Avar Khanate. The Genetic Origin of Huns, Avars, and Conquering Hungarians by Maróti et al. in 2022, confirms that only the elite preserved East-Asian “immigrant core” ancestry; most individuals plot on the European cline (EU-cline) with variable Pontic-Steppe and Caucasus admixture.

G-M406 did not arrive with the original Ugrian-speaking peoples or the Xiongnu-derived Avar elite. Instead, it reached the Avars through the same Northern Route mechanism: as a Caucasian/Pontic substrate carried by incorporated Sarmatian and Alan groups. These men were integrated into Avar society, buried with typical Avar grave goods, and appear in cemeteries across Hungary and Austria. However, they show no East-Asian autosomal signal. The same pattern continues into the Volga-Ural zone and the early Hungarian (Magyar) period.

Avar aDNA Samples
Sample:  RKF255
Location: Hungary
Date: AD 600-800
Y Haplogroup: G-FGC5089>M3302>TY500549

Sample:  RKF258
Location: Hungary
Date: AD 600-800
Y Haplogroup: G-FGC5089>M3302>TY500549

Sample: MGS488
Location: Mödling, Austria (South of Vienna)
Date:  AD 600-800 AD
Y haplogroup: FGC5089>FGC5081>G-Y2724>L14

Sample: KK1-245
Location: Kiskundorozsma, Hungary,
Date: AD 675–830; Late Avar
Y Haplogroup: G-FGC5089>FGC5081>L14>S19451>FGC5185>FGC5107>Z17085* 

Sample: AHPS209W
Location: Bodajk, Hungary
Date: AD 700-900
Y Haplogroup: G-M406>FGC5089>M3302>M3240>M3336>M3272

Sample: I33872
Location: Possibly Hungary
Date: AD 600-800
Y Haplogroup: G-M406>PF3299>FGC5089>M3302>M3425>M3422>Z36772
Status: Sample from unreleased paper Akbari2026

Sample: S47899
Location: Possibly Hungary
Date: AD 600-800
Y Haplogroup: G-M406>PF3299>FGC5089>M3302>BY21756>BY21758>BY21767>BY21760>M5843
Status: Sample from unreleased paper Akbari2026

Sample: I44285
Location: Possibly Hungary
Date: AD 600-800
Y Haplogroup: G-M406>PF3299>FGC5089>FGC5081>Y2724>FGC5124>FT49621
Status: Sample from unreleased paper Akbari2026

Sample: I47324
Location: Possibly Hungary
Date: AD 600-800
Y Haplogroup: G-M406>PF3299>FGC5089>FGC5081>Y2724>FGC5124>FT49621
Status: Sample from unreleased paper Akbari2026

Sample: I35428
Location: Possibly Hungary
Date: AD 600-800
Y Haplogroup: G-M406>PF3299>FGC5089>FGC5081>Y2724>FGC5124>L14>FGC5155>FGC5096>Z17084>FT45013>FTD7396
Status: Sample from unreleased paper Akbari2026

Sample: I28225
Location: Possibly Hungary
Date: AD 600-800
Y Haplogroup: G-M406>PF3299>PF3293>PF3296>Z30814>FGC41427>S11415
Status: Sample from unreleased paper Akbari2026

Post-Avar Hungary aDNA Samples
Sample: I33871
Location: Possibly Hungary
Date: AD 900-1100
Y Haplogroup: G-M406>PF3299>FGC5089>M3302>M3425>M3422>Z36772
Status: Sample from unreleased paper Akbari2026

Sample: I47353
Location: Possibly Hungary
Date: AD 700-900
Y Haplogroup: G-M406>PF3299>FGC5089>M3302>FT56871>BY21756>BY21758>BY21767>BY21760>M584
Status: Sample from unreleased paper Akbari2026

Sample: I34983
Location: Possibly Hungary
Date: AD 700-1100
Y Haplogroup: G-M406>PF3299>FGC5089>FGC5081>Y2724>Z17887>L645>FTA2742>PH1227>BY104556
Status: Sample from unreleased paper Akbari2026

The Russian Branches (Volga-Ural Corridor)

While the westward flow of G-M406 through the Avar Khaganate into the Carpathian Basin is now well documented, the same Northern Route simultaneously extended eastward along the Volga River and into the Volga-Ural region. 

Avar and Bulghar interactions with Khazar frontier groups allowed the lineage to reach the Volga-Ural zone (Novinki culture), where it mixed with proto-Magyars in the Karayakupovo Horizon. 

By the late 9th century, the Novinki group was absorbed into the rising Volga Bulgaria. This new state eventually dominated both the steppe nomads and many of the nearby Finno-Ugric tribes. The Volga Bulgars did not control the upper Volga, where a G-M406 sample was found amongst the Merya in Shekshovo, Russia dating to about AD 841. This spread and assimilation further up the Volga was likely facilitated by trade as Volga Bulgarian items such as ceramics and jewelry have been found in Shekshovo.

It was due to conflict between a separate branch of Bulgars from the Danube that the Magyars set out for Carpathain Basin. Following a Byzantine-sponsored Magyar invasion of Bulgaria, Simeon I of the Danube Bulgars retaliated by hiring the Pechenegs to strike the Magyar heartland while their warriors were away on campaign. This brutal "backstab" destroyed the Magyar settlements, leaving the returning army with no home to return to and Pecheneg enemies to their east and Bulgars to their south. With no other viable territory, Magyar Prince Árpád led his people across the Carpathian Mountains in AD 895 to conquer their modern-day homeland.

When the Magyars, known today as the Hungarians, conquered the Carpathian Basin, they carried a pre-existing Pontic substrate with them that contained men of G-M406. Upon arrival, they likely encountered a pre-existing substrate of G-M406 already seeded on the basin from prior nomadic groups. Two high-coverage Russian aDNA samples illustrate this beautifully: they show similar downstream G-M406 branches to those already seen among Scythians, Sarmatians, and Avars, yet they appear in distinctly non-Ugric and non-Xiongnu contexts. This shows the lineage was carried as a minority Caucasian/Pontic marker by Bulghar, Khazar, and Merya networks.

G-M406 samples with complete Avar-Hungary and Avar-Austria datasets

Possible Kushan Connection

The Scythian world was never limited to the Pontic Steppe. The eastern branch, the Saka (or Eastern Scythians). originated in the same Central Asian steppe zone that the western Scythians and Cimmerians had already reached by about 900 BC. Around 140–100 BC the Saka were pushed southward by the incoming Yuezhi and poured into the region of Bactria (modern Tajikistan, Uzbekistan, northern Afghanistan). This is exactly the region and timeframe of the Kushan sample I12293 from Tajikstan circa 200 BC - AD 100. These Saka groups were not a separate people, they shared language, horse-nomad culture, and genetic substrate with the Pontic Scythians. The same Caucasus/Anatolian-derived G-M3302 men who had already seeded the Pontic Steppe may have simply rode the eastern arm of the same nomadic wave.

According to the publishers of The Formation of Human Populations in South and Central Asia, the sample is autosomally anywhere between 20 to 37 percent N. Anatolian.  While the individual himself resided within the region historically associated with the Bactria-Margiana Archaeological Complex (BMAC), his genetic profile indicates a significant ancestral connection to Middle Bronze Age Armenians. This suggests that one of his immediate ancestors migrated from Armenia to the BMAC region. Furthermore, the individual's genome shows an additional genetic shift towards the local BMAC population, meaning that this migration likely occurred within the timeframe of a close ancestor rather than in the distant past. 

I12293's paternal line and recent Armenian ancestry may suggest that G-M3302 men continued to move out of the Caucasus/Anatolia reservoir during the height of Scythian/Saka expansion. The same branch already documented on the Pontic Steppe simply followed the eastern nomadic vector. Modern downstream M3302 testers in Pakistan and northern India may represent the final southern extension of this movement: Indo-Scythian (Saka) descendants who pushed even farther into the subcontinent in the 1st century BC–AD.

This sample therefore closes the eastern loop of the Northern Route: the identical M3302 lineage that seeded the Pontic Steppe (Cimmerian/Scythian) also radiated eastward with the Saka, reaching as far as the Kushan heartland and ultimately leaving traces in modern Pakistani populations. It is not a separate migration, it is the symmetrical eastern mirror of the western path that later reached the Avars, Hungarians, and Volga-Ural zone.

Kushan aDNA Sample
Sample:  I19086
Location: Ksirov, Tajikstan
Date: 100 BC - AD 300
Y Haplogroup: G-M406

Possible Northern Expansion of G-M406 illustrated by PCA Autosomal Shifts

• Neolithic & Chacolithic Anatolia

• Bronze Age Caucusus

• Iron Age Pontic Steppe

• Expansion west into Hungary

• Expansion up Volga River

• Indo-Scythian Expansion east into Bactra region