Scandinavian Hunter-Gatherers were identified as a distinct ancestral component in a study published in Nature in 2014. A number of remains examined at Motala, Sweden, and a separate group of remains from 5,000 year-old hunter-gatherers of the Pitted Ware culture, were identified as belonging to SHG. An SHG individual from Motala was identified as being of 81% Western Hunter-Gatherer ancestry, and 19% Ancient North Eurasian ancestry. In a genetic study published in Nature in March 2015, the remains of six SHGs buried at Motala between ca. 6000 BC and 5700 BC. Of the four males surveyed, three carried the paternal haplogroup I2a1 or various subclades of it, while the other carried I2c. With regards to mtDNA, four individuals carried subclades of U5a, while two carried U2e1. The study found SHGs to constitute one of the three main hunter-gatherer populations of Europe during the Holocene. The two other groups were WHGs and EHGs, between whom SHGs formed a distinct cluster. SHGs living between 6000 BC and 3000 BC were found to largely genetically homogenous, with little admixture occurring among them during this period. EHGs were found to be more closely related to SHGs than WHGs. In a genetic study published in Nature in November 2015, the six SHGs from Motala were subjected to further analysis. The study found it possible to model SHGs as a mixture of WHGs and EHGs. SHGs appeared to have persisted in Scandinavia until after 5,000 years ago. Results from studies of SHGs were found to be surprising. The Motala SHGs were found to be closely related to WHGs. In three samples, the haplotype carrying the derived allele of rs3827760 in the EDAR gene, which is today common in East Asia, but largely absent in modern-day Europe outside Scandinavia. This haplotype is not of East Asian origin however. Most of the Motala SHGs were found to have the depigmentation alleles SLC45A2 and SLC24A5. A genetic study published in Nature in July 2016 found SHGs to be a mix of EHGs and WHGs. WHGs were in turn a mix of EHGs and the Upper Paleolithic people of the Grotte du Bichon in Switzerland. EHGs derived 75% of their ancestry from ANEs. In a genetic study published in PLOS Biology in January 2018, the remains of seven SHGs were examined. All three samples of Y-DNA extracted belonged to subclades of I2. With respects to mtDNA, four samples belonged to U5a1 haplotypes, while three samples belonged U4a2 haplotypes. All samples from western and northern Scandinavia carried U5a1 haplotypes, while all the samples from eastern Scandinavia except from one carried U4a2 haplotypes. The authors of the study suggested that SHGs were descended from a WHG population that had entered Scandinavia from the south, and an EHG population which had entered Scandinavia from the northeast along the coast. The WHGs who entered Scandinavia are believed to have belonged to the Ahrensburg culture. These WHGs and EHGs had subsequently mixed, and the SHGs gradually developed their distinct character. The SHGs from western and northern Scandinavia had less more EHG ancestry than individuals from eastern Scandinavia. The SHGs were found to have a genetic adaption to high latitude environments, including high frequencies of low pigmentation variants and genes designed for adaption to the cold and physical performance. SHGs displayed a high frequency of the depigmentation alleles SLC45A2 and SLC24A5, and the OCA/Herc2, which effects eye pigmentation. These genes were much less common among WHGs and EHGs. A surprising continuity was displayed between SHGs and modern populations of Northern Europe in certain respects. Most notably, the presence of the protein TMEM131 among SHGs and modern Northern Europeans was detected. This protein may be involved in long-term adaptation to the cold. In a genetic study published in Nature Communications in January 2018, the remains of an SHG female at Motala, Sweden between 5750 BC and 5650 BC was analyzed. She was found to be carrying U5a2d and "substantial ANE ancestry". The study found that Mesolithic hunter-gatherers of the eastern Baltic also carried high frequencies of the HERC2, SLC45A2 and SLC24A5 alleles. They however harbored less EHG ancestry than SHGs. Genetic continuity between the SHGs and the Pitted Ware culture of the Neolithic was detected. The results further underpinned previous suggestion that SHGs were descended from northward migration of WHGs and a subsequent southward migration of EHGs. A certain degree of continuity between SHGs and northern Europeans was detected. A study published in Nature in February 2018 included an analysis of a large number of individuals of prehistoric Eastern Europe. Thirty-seven samples were collected from Mesolithic and Neolithic Ukraine. These were determined to be an intermediate between EHG and SHG. Samples of Y-DNA extracted from these individuals belonged exclusively to R haplotypes ) and I haplotypes. mtDNA belonged almost exclusively to U.
Physical appearance
Genomic data suggest that SHGs had darker skin than EHGs but lighter skin than WHGs and blue to light brown eye color. This is strikingly different from WHGs and EHGs, who are believed to have been blue eyed and dark skinned, and brown-eyed and light-skinned, respectively. On the basis of archaeological and genetic evidence, the Swedish archaeologist Odin Nilsson has made forensic reconstructions of both male and female SHGs.
