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Genetics

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Ancient DNA studies have conclusively demonstrated that agriculture was spread by migration,[1] and demic diffusion played a major role in the introduction of the Neolithic cultural package to Northern Europe.[2] In Scandinavia, the first evidence for farming is associated with the Funnelbeaker Culture (FBC).[3] FBC farmers were genetically similar to other Early Neolithic farming populations,[4] with predominantly Anatolian Farmer-related ancestry, but also with a relatively high Mesolithic Western Hunter-Gatherer component compared with other European farmer populations.[5][6] FBC individuals have been modelled as c. 80% Anatolian Farmer-related and c. 20% Western Hunter-Gatherer-related ancestry.[a][7][8] FBC populations have some of the largest amounts of Western Hunter-Gatherer admixture (c. 10-35%) found in any Neolithic Farmer population.[5]

FBC groups are genetically distinct from individuals associated with the partly contemporaneous Pitted Ware Culture (PWC) and those of the subsequent Battle Axe Culture; while Funnelbeaker individuals are genetically similar to other European farmer populations, Pitted Ware Culture individuals are closely related to earlier Scandinavian Hunter-Gatherers (SHG), and individuals found in Battle Axe Culture contexts are similar to other Corded Ware Culture individuals.[9]

Neolithic FBC individuals in Denmark form a genetic cluster with individuals from FBC contexts in Sweden and Poland,[5] and Middle Neolithic Polish FBC individuals from Kuyavia form a cluster with Lengyel, Linear Pottery, Anatolian, Hungarian, and Iberian farmers.[10] According to Sánchez-Quinto et al. 2018, Scandinavian FBC individuals are more closely related to British Neolithic individuals than they are to Central European farmers, perhaps as a result of migration between the British Isles and Scandinavia.[11] Polish FBC individuals are significantly more closely related to Scandinavian FBCs than they are to Baltic Early and Middle Neolithic individuals.[12] Like other Central European Neolithic Farmers, FBC farmers were genetically similar to modern Southern Europeans, especially Sardinians.[10]

Serrano et al. 2021 propose that the formation of the FBC population must be either the result of a WHG-like population admixing with Anatolian Farmers in Central Europe, or farmer populations in Scandinavia admixing with local WHG-like hunter-gatherers.[13] Scandinavian Hunter-Gathers, a population intermediate between WHG and EHG,[14] have been ruled out as a source of the hunter-gatherer component in FBC individuals.[15][13] According to Mittnik et al. 2018, the hunter-gatherer component in a FBC female from Southern Sweden came from either a WHG-like or Baltic Hunter-gatherer-like population.[15] Allentoft et al. 2024 conclude that the WHG component in Danish FBC-associated individuals was not derived from a local source,[5] and Papac et al. 2021 found that the ancestry of Bohemian Funnelbeaker individuals was mainly non-local, estimating that admixture with a WHG-like population occurred in the ancestors of the Bohemian Funnelbeaker population c. 5,079-4,748 BCE.[16] García-Puchol et al. 2017 propose that the ancestors of the FBC population probably admixed with Mesolithic hunter-gatherers before expanding.[4] According to Allentoft et al. 2024, FBC farmers in Denmark are closely related to Polish Globular Amphora Culture (GAC) individuals, suggesting that the genetic origins of the FBC were in Eastern Europe.[5]

In some regions, such as Denmark, additional limited admixture between FBC groups and Mesolithic hunter-gatherer populations occurred after c. 5,400 BP. In Sweden, according to Allentoft et al. 2024, no admixture between FBC individuals and local hunter-gatherers was detected.[5] However, according to Sánchez-Quinto et al. 2019, hunter-gatherer admixture is found on both the autosomes and X-chromosome in FBC individuals from the Ansarve and Gökhem sites in Sweden, which they suggest supports a scenario of recent admixture with local Northern European hunter-gatherers.[17]

Although Mittnik et al. 2018 suggest that PWC individuals can feasibly be modelled as an admixture of Scandinavian Hunter-Gatherers (c. 74%) and Early Neolithic FBC (c. 26%),[15] other studies have detected only minimal gene flow between FBC and PWC or SHG groups, even though hunter-gatherer groups in Scandinavia lived in close proximity with FBC individuals for over 1000 years.[13][18]

The population associated with the Battle Axe Culture, a regional Corded Ware Culture (CWC) group which replaced the FBC in southern Scandinavia, may be the result of admixture between migrating CWC groups from the southern Baltic, who carried high levels of Yamnaya-related ancestry, and FBC groups in Scandinavia. Malmström et al. 2019 suggest that a model with proportions of 32% FBC and 68% CWC related ancestries is feasible.[19] In Denmark, in the transition to the Single Grave Culture, the Neolithic Farmer population were substantially replaced from c. 4,800 BP by migrating populations carrying high levels of Yamnaya ancestry, with minimal contribution from local Scandinavian farmers. Allentoft et al. 2024 propose that the SGC can be modelled as c. 60–85% Yamnaya-related, c. 10–23% Eastern European GAC, and c. 3–18% Scandinavian FBC.[20]

Published Y-DNA haplogroups from Funnelbeaker-associated males include I2a1b1a1 (Esperstedt, Salzmünde/Bernberg culture), I (Baalberge culture), R1b1a (Baalberge culture), G2a2a (Salzmünde-Schiebzig),[21][22][23] IJK (x J) (Salzmünde-Schiebzig),[24] and IJ-M429 (Rössberga).[25] <---SOME OF THESE SAMPLES ARE TERRIBLE COVERAGE

Four males from a megalithic burial at Ansarve, Gotland were assigned to haplogroups I2a1b1a1, I2a1b1a1, I2a1b, and I2a1b1a1.[17] The I2 Y-DNA haplogroup of the Ansarve individuals is common among European Mesolithic hunter-gatherers, but not in European Early Farmer groups. It is frequently found in other 4th millenium BCE farming cultures. The preponderance of the I2a male lineage at the Ansarve site may suggest male-biased admixture between farmers and hunter-gatherers over time. The uniformity of Y-DNA lineages along with over-representation of males and an identified second degree family relationship in the Ansarve tomb imply that the Ansarve individuals and other megalith building communities were part of a patrilineal and stratified society.[26]

Although the high proportion of I2a male lineages found in individuals associated with the FBC and other megalithic societies suggests that large numbers of hunter-gatherer males mixed with 4th millenium BCE farming groups, comparison of affinity to Mesolithic hunter gathers on the autosomes and X-chromosome of the Ansarve and Gökhem individuals suggests that sex-biased admixture occurred mostly between male farmers and female hunter-gatherers, according to Sánchez-Quinto et al. 2019.[27]

The mtDNA haplogroup frequencies from the Middle Neolithic Salzmünde and Bernburg groups (Germany) published in Haak et al. 2015 are T2 (30%), J (20%), U5b (20%), K (10%), H (10%), and U5a (10%).[28] The mtDNA frequencies for the Funnelbeaker culture published in Juras et al. 2021 are separated into North West (Scandanavian and Western European) and Central East (Poland) groups.[29] For the North West group, the frequencies are H (36.842%) T (21.053%), J (15.789%), U5b (10.526%), U3 (5.263%), U5a (5.263%), and K (5.263%). For the Central East group, the frequencies are H (34.615%), U5b (15.385%), T (11.538%), W (11.538%), X (7.692%), J (3.846%), U (3.846%), U3(3.846%), U5a (3.846%), and K (3.846%). In individuals from passage graves at Västergötland and Öland (Sweden), mtDNA haplogroups K, T, H, J1 and J2 have been found.[30]

Mitochondrial DNA haplogroups from FBC groups are a mixture of a relatively high proportion of hunter-gatherer lineages (such as U5a, U5b) as well as Neolithic Farmer lineages (e.g. T2, J, K). Brandt et al. 2013 gives figures of c. 30% hunter-gatherer and c. 60% Neolithic Farmer mtDNA haplogroups in Scandinavian FBC individuals, values which are intermediate between hunter-gatherer groups and the Early to Middle Neolithic individuals from the Mittelelbe-Saale region.[31] Brandt et al. 2015 suggests figures of 20% hunter-gatherer and 40% Neolithic Farmer maternal lineages.[32] Juras et al. 2021 found that the mtDNA haplogroups of Central European FBCs were mainly of Neolithic farmer origin. However haplogroup U5b, associated with Western Hunter-Gatherers, was the second most common mtDNA lineage (more than 15% of the total) among Central East (Poland) FBCs.[33] Hunter-gatherer mtDNA lineages are also found in individuals associated with the Gökhem group.[30]

At Scandanavian FBC megalithic sites in Ansarve, Gotland, mtDNA haplogroup diversity was greater than Y-DNA haplogroup diversity. The distribution of uniparental haplogroups at these sites was similar to that of other 4th millennium BCE Neolithic Farmers. [34][35] The mtDNA diversity at Ansarve shows that this burial site was not dominated by a single maternal lineage.[36][37]

Similar mtDNA haplogroup distribution to that observed among groups from Swedish passage graves, such as higher proportions of haplogroup H, is found in individuals from other Central European Early to Middle Neolithic collective burials. The haplogroup distribution also shows affinity to the Early Neolithic Gurgy group from the Paris Basin and Southwest European Neolithic populations, suggesting complex interactions between Middle Neolithic European groups.[38] Later Central European FBC groups of the Mittelelbe-Saale region, such as the Bernburg culture, had higher proportions of hunter-gatherer (29.4%) and lower proportions of farmer (47.1%) mtDNA lineages compared to preceding Neolithic populations in the same area, similar figures to the Scandinavian FBC groups.[31] Increased frequencies of haplogroups U5a and U5b were found in the Bernburg culture, suggesting that a migration of Funnelbeaker groups from the north to Central Europe occurred c. 3100 BCE. [39] Regional differences in mtDNA haplogroup distribution among various FBC groups suggest different migration patterns and admixture events in different areas of the Funnelbeaker Culture.[37]

Juras et al. 2021 found that FBC populations were maternally related to individuals from th3 Złota culture and other GAC groups. They found no maternal relationship between FBC individuals and CWC groups [40]

Admixture between FBC and PWC groups is suggested by the presence of mtDNA haplogroups J and K in both groups, according to García-Puchol et al. 2017[4]

Notes

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  1. ^ According to Allentoft et al. 2024, the Anatolian Farmer-related ancestry in FBC individuals averages around 79%.[7] According to Papac et al. 2021, FBC individuals from Bohemia could on average be modelled as c. 80.4% Anatolian Farmer-related and c. 19.6% WHG-related.[8]

Citations

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  1. ^ Whittle et al. 2023, p. 41.
  2. ^ Malmström et al. 2015, p. 8.
  3. ^ Malmström et al. 2015, p. 1.
  4. ^ a b c García-Puchol et al. 2017, p. 328.
  5. ^ a b c d e f Allentoft et al. 2024, p. 333.
  6. ^ Juras et al. 2021, p. 3.
  7. ^ a b Allentoft et al. 2024, Supplementary Data 4.
  8. ^ a b Papac et al. 2021, Supplementary Table S12.
  9. ^ Malmström et al. 2020, pp. 645–646.
  10. ^ a b Fernandez et al. 2018, p. 2.
  11. ^ Sánchez-Quinto et al. 2019, p. 9470,9473.
  12. ^ Fernandez et al. 2018, p. 3.
  13. ^ a b c Serrano et al. 2021, p. 182.
  14. ^ Mittnik et al. 2018, p. 2.
  15. ^ a b c Mittnik et al. 2018, p. 6.
  16. ^ Papac et al. 2021, p. 4.
  17. ^ a b Sánchez-Quinto et al. 2019, p. 9471.
  18. ^ Malmström et al. 2020, p. 639.
  19. ^ Malmström et al. 2019, pp. 5–6.
  20. ^ Allentoft et al. 2024, p. 334.
  21. ^ Haak et al., 2015 & Table 1, Rows 38, 159, 112, 116.
  22. ^ Mathieson et al. 2018, Supplementary Table 1, Row 126-130.
  23. ^ Lipson et al. 2017, Sup Table 1, Row 118.
  24. ^ Lipson et al. 2017, Sup Table 1, Row 120.
  25. ^ Malmström et al. 2019, p. 3.
  26. ^ Sánchez-Quinto et al. 2019.
  27. ^ Sánchez-Quinto et al. 2019, pp. 66–68, Supplement.
  28. ^ Haak et al. 2015, p. 19, Supplementary Information.
  29. ^ Juras et al. 2021, Appendix S2, Table S2.
  30. ^ a b Fraser et al. 2018, p. 326.
  31. ^ a b Brandt et al. 2013, p. 260.
  32. ^ Brandt et al. 2015, p. 10.
  33. ^ Juras et al. 2021, p. 12.
  34. ^ Sánchez-Quinto et al. 2019, p. 9470.
  35. ^ Sánchez-Quinto et al. 2019, p. 67, Supplement.
  36. ^ Malmström et al. 2015, p. 7.
  37. ^ a b Fraser et al. 2018, p. 332.
  38. ^ Fraser et al. 2018, p. 331.
  39. ^ Brandt et al. 2015, p. 11.
  40. ^ Juras et al. 2021, p. 11.

Bibliography

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  • Allentoft, Morten E.; Sikora, Martin; Fischer, Anders; Sjögren, Karl-Göran; Ingason, Andrés; Macleod, Ruairidh; Rosengren, Anders; Schulz Paulsson, Bettina; Jørkov, Marie Louise Schjellerup; Novosolov, Maria; Stenderup, Jesper; Price, T. Douglas; Fischer Mortensen, Morten; Nielsen, Anne Birgitte; Ulfeldt Hede, Mikkel (2024-01-11). "100 ancient genomes show repeated population turnovers in Neolithic Denmark". Nature. 625 (7994): 329–337. doi:10.1038/s41586-023-06862-3. ISSN 0028-0836. PMC 10781617. PMID 38200294.{{cite journal}}: CS1 maint: PMC format (link)
  • Whittle, Alasdair; Pollard, Joshua; Greaney, Susan (2023). Ancient DNA and the European Neolithic: relations and descent. Neolithic Studies Group seminar papers. Oxford: Oxbow Books. ISBN 978-1-78925-910-0.
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