mtDNA haplogroup frequency analysis from Verteba Cave supports a strong cultural frontier between farmers and hunter-gatherers in the North Pontic steppe

New preprint paper at BioRxiv, led by a Japanese researcher, with analysis of mtDNA of Trypillians from Verteba Cave, Analysis of ancient human mitochondrial DNA from Verteba Cave, Ukraine: insights into the origins and expansions of the Late Neolithic-Chalcolithic Cututeni-Tripolye Culture, by Wakabayashi et al. (2017).

Abstract:

Background: The Eneolithic (~5,500 yrBP) site of Verteba Cave in Western Ukraine contains the largest collection of human skeletal remains associated with the archaeological Cucuteni-Tripolye Culture. Their subsistence economy is based largely on agro-pastoralism and had some of the largest and most dense settlement sites during the Middle Neolithic in all of Europe. To help understand the evolutionary history of the Tripolye people, we performed mtDNA analyses on ancient human remains excavated from several chambers within the cave.

Results: Burials at Verteba Cave are largely commingled and secondary in nature. A total of 68 individual bone specimens were analyzed. Most of these specimens were found in association with well-defined Tripolye artifacts. We determined 28 mtDNA D-Loop (368 bp) sequences and defined 8 sequence types, belonging to haplogroups H, HV, W, K, and T. These results do not suggest continuity with local pre-Eneolithic peoples, but rather complete population replacement. We constructed maximum parsimonious networks from the data and generated population genetic statistics. Nucleotide diversity (π) is low among all sequence types and our network analysis indicates highly similar mtDNA sequence types for samples in chamber G3. Using different sample sizes due to the uncertainly in number of individuals (11, 28, or 15), we found Tajima’s D statistic to vary. When all sequence types are included (11 or 28), we do not find a trend for demographic expansion (negative but not significantly different from zero); however, when only samples from Site 7 (peak occupation) are included, we find a significantly negative value, indicative of demographic expansion.

Conclusions: Our results suggest individuals buried at Verteba Cave had overall low mtDNA diversity, most likely due to increased conflict among sedentary farmers and nomadic pastoralists to the East and North. Early Farmers tend to show demographic expansion. We find different signatures of demographic expansion for the Tripolye people that may be caused by existing population structure or the spatiotemporal nature of ancient data. Regardless, peoples of the Tripolye Culture are more closely related to early European farmers and lack genetic continuity with Mesolithic hunter-gatherers or pre-Eneolithic groups in Ukraine.

Genetic finds keep supporting the long-lasting cultural and linguistic frontier that Anthony (2007) – among others – asserted existed in the North-West Pontic steppe in the Mesolithic and Neolithic, between western steppe cultures and farmers, while it disproves Kristiansen’s theories of Sredni Stog expansion in Kurgan waves with a mixture of GAC and Trypillia within the Corded Ware culture:

Previous ancient DNA studies showed that hunter-gatherers before 6,500 yrBP in Europe commonly had haplogroups U, U4, U5, and H, whereas hunter-gatherers after 6,500 yrBP in Europe had less frequency of haplogroup H than before. Haplogroups T and K appeared in hunter-gatherers only after 6,500 yrBP, indicating a degree of admixture in some places between farmers and hunter-gatherers. Farmers before and after 6,500 yrBP in Europe had haplogroups W, HV*, H, T, K, and these are also found in individuals buried at Verteba Cave. Therefore, our data point to a common ancestry with early European farmers. Our data also suggest population replacement. Mathieson et al. analyzed a number of Neolithic Ukrainian samples (petrous bone) from several sites in southern, northern, and western Ukraine, dating to ~8,500 – 6,000 yrBP, and found exclusively U (U4 and U5) mtDNA lineages. It should be noted that ‘Neolithic’ in this context does not mean the adoption of agriculture, but rather simply coinciding with a change in material culture. They also analyzed several Trypillian individuals from Verteba Cave (different samples from the those included in this study). Similar to our findings, they found a wider diversity of mtDNA lineages, including H, HV, and T2b. These data, combined with our results, appear to confirm almost complete population replacement by individuals associated with the Tripolye Culture during the Middle to Late Neolithic.

The findings also hint to potential contacts of Yamna with Usatovo as predicted by Anthony (2007), or alternatively (lacking precise dates) to contacts with Corded Ware migrants:

Trypillians were very much a distinct people who most likely displaced 1 local hunter-gatherers with little admixture. Haplogroup W was also observed in several specimens deriving from Site G3. Although we are unsure if all of these haplogroups come from a single or multiple individuals, this observation is interesting in that it is relatively rare and isolated among Neolithic samples. It has, however, been found in samples dating to the Bronze Age. In the study by Wilde et al. [35], they found haplogroup W present in two samples from the Early Bronze Age associated with the Yamnaya and Usatovo cultures. The Usatovo culture (~ 3500 – 2500 BC) was found in Romania, Moldova, and southern Ukraine. It was the conglomeration of Tripolye and North Pontic steppe cultures. Therefore, this individual could link the Trypillian peoples to the Usatovo peoples and perhaps to the greater Yamnaya steppe migrations during the Bronze Age that lead to the Corded Ware Culture.

On the other hand, an article written in terms of mtDNA haplogroup frequencies seems to offer too little proof of anything today. The lack of Y-DNA haplogroups and data on admixture makes their interpretations provisional, subject to change when these further data are published. Also, radiocarbon dating is only confident for individuals of one site (site 7), dated ca. 5,500 cal BP, while “other chambers in the cave are not as confidently dated”…

verteba-cave-mtDNA
“Based on the 8 sequence types of the mtDNA D-loop, a maximum parsimonious phylogenetic network was constructed. Circles represent the sequence types, and the size of the circle is proportional to the number of samples. Numbers on the branches between the circles are nucleotide position numbers (+16,000) of the human mitochondrial genome sequence (rCRS). Information about the location (chamber within the cave) where the specimen was excavated is also provided. Areas 2 and 17 are part of Site 7, and these are defined as a separate chamber, although they are located in close proximity within Site 7. The other chambers, Site 20, G2, and G3, are independent and separate locations within the cave. ‘Undefined’ chamber describes an unknown location within the cave. Specimens from each chamber showed deviation for the sequence type distribution observed in the sample set. For example, specimens excavated from Site 7 had five unique sequence types, (I, II, III, IV, and VIII), while specimens excavated from chamber G 21 had mainly one sequence type (V)”. Made available by the authors under a CC-BY-NC-ND 4.0 International license.

We had also seen signs of conflict between Trypillian and steppe cultures in a recent article, Violence at Verteba Cave, Ukraine: New Insights into the Late Neolithic Intergroup Conflict, by Madden et al. (2017):

Many researchers have pointed to the huge “megasites” and construction of fortifications as evidence of intergroup hostilities among the Late Neolithic Tripolye archaeological culture. However, to date, very few skeletal remains have been analyzed for the types of traumatic injury that serve as direct evidence for violent conflict. In this study, we examine trauma on human remains from the Tripolye site of Verteba Cave in western Ukraine. The remains of 36 individuals, including 25 crania, were buried in the gypsum cave as secondary interments. The frequency of cranial trauma is 30-44% among the 25 crania, six males, four females and one adult of indeterminate sex displayed cranial trauma. Of the 18 total fractures, 10 were significantly large and penetrating suggesting lethal force. Over half of the trauma is located on the posterior aspect of the crania, suggesting the victims were attacked from behind. Sixteen of the fractures observed were perimortem and two were antemortem. The distribution and characteristics of the fractures suggest that some of the Tripolye individuals buried at Verteba Cave were victims of a lethal surprise attack. Resources were limited due to population growth and migration, leading to conflict over resource access. It is hypothesized that during this time of change burial in this cave aided in development of identity and ownership of the local territory.

Related:

Correlation does not mean causation: the damage of the ‘Yamnaya ancestral component’, and the ‘Future American’ hypothesis

New Ukraine Eneolithic sample from late Sredni Stog, near homeland of the Corded Ware culture

The concept of “outlier” in studies of Human Ancestry, and the Corded Ware outlier from Esperstedt

Marija Gimbutas and the expansion of the “Kurgan people” based on tumulus-building cultures

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