Ancient DNA studies of geographically and chronologically disparate human population groups are of the utmost importance for understanding their development and interactions which led eventually to the contemporary picture of human diversity and resettlement. Despite significant progress and accumulation of genomic data for ancient archaeological cultures in Eurasia 1–4, detailed data describing the genesis and evolution of populations associated with single archaeological culture and their sequences in certain historical regions are still insufficient 5–7.
Anatomically modern humans have inhabited the semi-arid steppe and forest-steppe regions of Southern Siberia since the Late Paleolithic period 8. During the Chalcolithic, the Bronze and Early Iron Age, this region became home to several human populations who greatly influenced the cultural and technological development and exchange processes in the middle latitudes of Eurasia and left a rich archaeological heritage 9.
The Minusinsk Basin is a mountainous-steppe area in the northern part of the Altai-Sayan plateau, which is situated at the juncture of the Siberian taiga and the mountains and steppe of Central Asia, on both sides of the middle reaches of the Yenisei River. This region corresponds to the modern territories of the Republic of Khakassia and the southern part of the Krasnoyarsk region in Russia. The Minusinsk Basin’s ancient human history had much in common with other ancient societies of the Altai-Sayan plateau but stood out from them due to its milder climate and less accessible location. The Minusinsk Basin facilitated a complex economy and relatively safe life for many ancient human societies because of its favorable conditions and abundance of natural resources 10.
The Minusinsk Basin was a place where different archaeological cultures with complex ideologies, advanced technologies and distinctive artistic activity flourished. This region is world-famous for its rich and elaborate rock art of various epochs, e.g. the monumental steles of the Okunev culture 9,11−14 or bronze artefacts of the Tagar culture 15,16. The Tes’ and early Tashtyk cultures in the later historical period are also well-known on account of their elaborate funeral customs including mummification, recreating the bodies, head modification, and face masks (Fig. 1B; Fig. 1C) 17–19. Interactions between archaeological cultures from adjacent and remote regions, including the Eastern European steppe, Central Asia, and the Chinese plain have been reliably attested by parallels in the local material culture, arts and ritual traditions since the Chalcolithic period 12.
The Minusinsk Basin has always been a well-populated area, leading to a very high density of archaeological sites 20,21. Many of them are still clearly visible in the steppe due to an age-old tradition of creating enclosures or other above-ground constructions from large stones and slabs. Notably, the Minusinsk Basin became the first region in Russia where archaeological excavations were carried out –as far back as 1722 22. The first periodization of antiquities from the Early Bronze to the Middle Ages was established in the 1920s based on local material culture 20, laying a reliable foundation for the modern chronological system of archaeological sites/artefacts of the Eurasian steppe. Moreover, this system is still in use for the study of the archaeology of the Eurasian steppe. The material culture, art and burial rituals of the ancient societies in this part of Southern Siberia have been extensively studied whereas the human population history is still somewhat sparsely established for different historical periods 12.
Patterns of interaction between migrant and local human populations changed over time in Southern Siberia23. Recent studies, which combined genetical, archaeological and physical anthropological data, have shown that the exchange of local populations caused by migrations into the Minusinsk Basin was almost total during the Chalcolithic Period and the Early Bronze Age. In the Late Bronze Age (LBA), however, the character of interaction between local and incoming groups changed, and more genetic and cultural continuity of ancient societies can be observed from the mid-2nd millennium BC 12. Extensive archaeological data from the period between the LBA (represented by several archaeological horizons previously known as the Karasuk culture) and the Early Iron Age (Tagar and Tes’ cultures) suggest several local migrations into this region, and archaeological materials indicate both local and outside ancestry for each of the corresponding chronological horizons. Genetic information is very scarce for them, and nuclear genomic data from the LBA and Early Iron Age archaeological cultures of the Minusinsk Basin are of utmost importance for building the overall picture of the genomic ancestry in this region.
The Tashtyk culture is the post-Scythian Iron Age archaeological culture of the Minusinsk Basin which followed the Tagar culture 18. It is viewed as having had a nomadic pastoralist economy supplemented by hunting-gathering and small-scale agriculture. Cultural innovations and a slight admixture of East Asian craniological features on early Tashtyk skulls and funeral masks led to the idea of some mass migration from Central Asia which possibly took place at the turn of the Christian era 24–26. This East Asian craniological impact is now seen as considerably exaggerated for the early Tashtyk population 27 and requires further study. The interactions between local and various incoming human groups at that time are currently seen as a complex phenomenon, suggesting their co-existence in Southern Siberia during this historical period 28,29. Significant innovations documented in the burial rituals and grave goods of the early Tashtyk sites are still considered to be the result of migrations from Central Asia, likely from the areas adjacent to the Tarim Basin, but the question remains open 30.
To date, only a few studies on the genetic ancestry of the early Tashtyk culture have been carried out using Y-chromosome short tandem repeat (STR) analysis and mtDNA hypervariable region 1 (HVR1) sequencing. A study of 4 female and 1 male individuals from the Abakano-Pérévoz I cemetery provided information on their mitochondrial DNA (mtDNA) and Y-chromosome haplogroups31. C, HV, H, N9a, and T1 mtDNA and R1a Y-chromosome haplogroups were described for these individuals31.
The Oglakhty cemetery is one of the well-studied Tashtyk burial complexes, situated 40 km north of the city of Abakan (Fig. 1A). This archaeological site with well-preserved organic artefacts and anthropological remains provides exceptionally detailed information on funeral rituals and peculiarities of material culture, which is usually lost. People were buried in the Oglakhty graves according to different rites – mummification and cremation with the cremated remains (cremains) being placed into full-size human-like mannequins made from leather and grass 30,32,33. The graves are thought to be family ones as opposed to the previous and following periods’ collective burials containing tens or hundreds of dead. The biritualism of the early Tashtyk graves is a unique feature and possible evidence of the complex structure of this society made up of people with different origins and traditions.
Methods of archaeology and craniometry have up to now been unable to trace the core region, cultural and biological characteristics of the incoming human group that was incorporated into the early Tashtyk society. The family kinship of the dead from the Oglakhty graves cannot be established by these methods either, nor can the continuity of the early Tashtyk population from the previous local groups. Genetic studies of the individuals from the Oglakhty cemetery are capable of shedding light on these crucial questions.
Here we present nuclear genomes of two human mummies of the early Tashtyk culture (1st − 4th centuries AD) buried in the Oglakhty cemetery, grave 4 (Fig. 1D). A child mummy from the same grave has also been analyzed but its endogenous DNA content proved insufficient for cost-effective DNA sequencing, so we cannot take that information into consideration. In addition, the ancient DNA identification of animal leather and fur found in grave 4 in elements of the clothing and a mannequin was covered in this study.