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Scandinavia spanning the Iron Age to the present, based on 48 new and 249 published ancient genomes and genotypes from 16,638 modern individuals. We find regional variation in the timing and magnitude of gene flow from three sources: the eastern Baltic, the British-Irish Isles, and southern Europe. British-Irish ancestry was widespread in Scandinavia from the Viking period, whereas eastern Baltic ancestry is more localized to Gotland and central Sweden. In some regions, a drop in current levels of external ancestry suggests that ancient immigrants contributed proportionately less to the modern Scandinavian gene pool than indicated by the ancestry of genomes from the Viking and Medieval periods. Finally, we show that a north-south genetic cline that characterizes modern Scandinavians is mainly due to the differential levels of Uralic ancestry and that this cline existed in the Viking Age and possibly earlier. The authors acknowledge support from the National Genomics Infrastructure in Stockholm funded by Science for Life Laboratory, the Knut and Alice Wallenberg Foundation and the Swedish Research Council, and SNIC/Uppsala Multidisciplinary Center for Advanced Computational Science for assistance with massively parallel sequencing and access to the UPPMAX computational infrastructure. We used resources from projects SNIC 2022/23-132, SNIC 2022/22-117, SNIC 2022/23-163, SNIC 2022/22-299, and SNIC 2021-2-17. This research was supported by the Swedish Research Council project ID 2019-00849_VR and ATLAS (Riksbankens Jubileumsfond). Part of the modern dataset was supported by a research grant from Science Foundation Ireland (SFI), grant number 16/RC/3948, and co-funded under the European Regional Development Fund and by FutureNeuro industry partners.

Anatolia and the Near East have long been recognized as the epicenter of the Neolithic expansion through archaeological evidence. Recent archaeogenetic studies on Neolithic European human remains have shown that the Neolithic expansion in Europe was driven westward and northward by migration from a supposed Near Eastern origin [1-5]. However, this expansion and the establishment of numerous culture complexes in the Aegean and Balkans did not occur until 8,500 before present (BP), over 2,000 years after the initial settlements in the Neolithic core area [6-9]. We present ancient genome-wide sequence data from 6,700-year-old human remains excavated from a Neolithic context in Kumtepe, located in northwestern Anatolia near the well-known (and younger) site Troy [10]. Kumtepe is one of the settlements that emerged around 7,000 BP, after the initial expansion wave brought Neolithic practices to Europe. We show that this individual displays genetic similarities to the early European Neolithic gene pool and modern-day Sardinians, as well as a genetic affinity to modern-day populations from the Near East and the Caucasus. Furthermore, modern-day Anatolians carry signatures of several admixture events from different populations that have diluted this early Neolithic farmer component, explaining why modern-day Sardinian populations, instead of modern-day Anatolian populations, are genetically more similar to the people that drove the Neolithic expansion into Europe. Anatolia's central geographic location appears to have served as a connecting point, allowing a complex contact network with other areas of the Near East and Europe throughout, and after, the Neolithic.

The impact of human mobility on the northern European urban populations during the Viking and Early Middle Ages and its repercussions in Scandinavia itself are still largely unexplored. Our study of the demographics in the final phase of the Viking era is the first comprehensive multidisciplinary investigation that includes genetics, isotopes, archaeology, and osteology on a larger scale. This early Christian dataset is particularly important as the earlier common pagan burial tradition during the Iron Age was cremation, hindering large-scale DNA analyses. We present genome-wide sequence data from 23 individuals from the 10

Summary The processes leading up to species extinctions are typically characterized by prolonged declines in population size and geographic distribution, followed by a phase in which populations are very small and may be subject to intrinsic threats, including loss of genetic diversity and inbreeding [1]. However, whether such genetic factors have had an impact on species prior to their extinction is unclear [2, 3]; examining this would require a detailed reconstruction of a species' demographic history as well as changes in genome-wide diversity leading up to its extinction. Here, we present high-quality complete genome sequences from two woolly mammoths ( Mammuthus primigenius ). The first mammoth was sequenced at 17.1-fold coverage and dates to ∼4,300 years before present, representing one of the last surviving individuals on Wrangel Island. The second mammoth, sequenced at 11.2-fold coverage, was obtained from an ∼44,800-year-old specimen from the Late Pleistocene population in northeastern Siberia. The demographic trajectories inferred from the two genomes are qualitatively similar and reveal a population bottleneck during the Middle or Early Pleistocene, and a more recent severe decline in the ancestors of the Wrangel mammoth at the end of the last glaciation. A comparison of the two genomes shows that the Wrangel mammoth has a 20% reduction in heterozygosity as well as a 28-fold increase in the fraction of the genome that comprises runs of homozygosity. We conclude that the population on Wrangel Island, which was the last surviving woolly mammoth population, was subject to reduced genetic diversity shortly before it became extinct.

Scandinavia was one of the last geographic areas in Europe to become habitable for humans after the last glaciation. However, the origin(s) of the first colonizers and their migration routes remain unclear. We sequenced the genomes, up to 57x coverage, of seven hunter-gatherers excavated across Scandinavia and dated to 9,500-6,000 years before present. Surprisingly, among the Scandinavian Mesolithic individuals, the genetic data display an east-west genetic gradient that opposes the pattern seen in other parts of Mesolithic Europe. This result suggests that Scandinavia was initially colonized following two different routes: one from the south, the other from the northeast. The latter followed the ice-free Norwegian north Atlantic coast, along which novel and advanced pressure-blade stone-tool techniques may have spread. These two groups met and mixed in Scandinavia, creating a genetically diverse population, which shows patterns of genetic adaptation to high latitude environments. These adaptations include high frequencies of low pigmentation variants and a gene-region associated with physical performance, which shows strong continuity into modern-day northern Europeans.

This study deals with genetic analyses of an assemblage of mediaeval (13th century) cattle metapodials from Bern that had previously been osteometrically examined regarding sex, shape and wither height. The results from the genetic sexing of these small (height 100 to 120 cm) cattle correlate well with the osteometric interpretations. Some few exceptions we interpreted as cows used as draft animals with stouter bones and thus osteometrically determined as males. Two morphologically different groups of cow metatarsals however, we took as proof of the historical fact that Bern relied on livestock from different geographical origins: the town's vicinity and the alpine pastures with their favourable grazing conditions. It was not possible to distinguish them genetically. An analysis of one single nucleotide polymorphism (SNP) in the melanocortin receptor 1 (MC1R) showed that predominant coat colour most likely was red-brown. Furthermore, an analysis of the SNP in the Y-chromosomal intron UTY19 that divide modern taurine cattle in two major haplogroups (Y1 and Y2) showed that the mediaeval cattle belonged to the haplogroup Y2 with one single exception of a Y1.In der vorliegenden Arbeit wird über die Genom-Analyse einer Sammlung von Metapodien mittelalterlicher Hausrinder des 13. Jahrhunderts aus Bern berichtet. Geschlecht wie auch Widerristhöhe der zugrundeliegenden Tiere sind zuvor nach osteometrischen und morphologischen Kriterien bestimmt worden. Der genetische Befund der Geschlechtsbestimmung dieser mit einer Widerristhöhe von 100 bis 120 cm als klein befundenen Tieren korreliert gut mit demjenigen der Osteometrie. Davon ausgenommen sind einzelne Metacarpen, die morphologisch als männlich, genetisch jedoch als weiblich bestimmt wurden. Diese zählen wir wegen ihrer kräftigen Morphologie zu den als Zugtiere verwendeten Kühen. Das Vorhandensein von zwei morphologisch unterschiedlichen Gruppen von Kuh-Metatarsen erachten wir als Beleg dafür, dass Bern mit Schlachtvieh verschiedener geografischer Herkunft beliefert wurde: einerseits aus der Umgebung der Stadt und andererseits aus den Alpen, wo günstigere Futterbedingung herrschten. Es war nicht möglich, die beiden Gruppen genetisch zu unterscheiden. Die Untersuchung des Einzelnukleotid-Polymorphismus (SNP) im Melanocortinrezeptor 1 (MC1R) ergab für die Fellfarbe vorwiegend rot-braun. Weiter zeigte die Untersuchung des Y-chromosomalen SNPs im Intron des UTY19-Gens, nach welchem heutige taurine Rinder in die Haplogruppen Y1 und Y2 eingeteilt werden, dass die untersuchten mittelalterlichen Rinder mit einer Ausnahme (Y1) zur Haplogruppe Y2 gehören.

Hunters and Farmers The Neolithic period in Europe saw the transition from a hunter-gatherer lifestyle to farming. Previous genetic analyses have suggested that hunter-gatherers were replaced by immigrant farmers. Skoglund et al. (p. 747 , published online 24 April) sequenced one Mesolithic and nine Neolithic Swedish individuals to examine the transition from hunter-gatherers to farmers. Substantial genetic differentiation was observed between hunter-gatherers and farmers: There was lower genetic diversity within the hunter-gatherers and gene flow from the hunter-gatherers into the farmers but not vice versa.