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Distribution of Mesolithic and Neolithic Y-chromosome lineages, and their Bronze Age descendants. (A) Britain and Ireland with (B) zoom in on Orkney. Colors represent different Y-chromosome lineages, and distinct outlines represent the time period of the sample. Each circle represents one individual, except for Trumpington Meadows, Cambridgeshire (66), where two brothers are represented by a single circle. Maps prepared with GADM tools (https://gadm.org/data.html) (67) using data from SRTM (68).
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The Orcadian Neolithic has been intensively studied and celebrated as a major center of cultural innovation, whereas the Bronze Age is less well known and often regarded as a time of stagnation and insularity. Here, we analyze ancient genomes from the Orcadian Bronze Age in the context of the variation in Neolithic Orkney and Bronze Ag...
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... predominance of I2a1b-M423 is surprising because it is completely absent elsewhere in CA/BA Europe, where the Y-DNA landscape is heavily dominated by R1b-M269 (Figs. 2- (Figs. 3 and 4). Even in Britain and Ireland, outside of Orkney most Neolithic Y-DNA lineages belong to haplogroup I2a2-S33 or I2a2a-M223 (Fig. 3), although, curiously, our Neolithic individual from Skye belongs to the very rare I2a2b-S154, seen elsewhere only in Middle Neolithic France (43). I2a1b-M423 seems to be largely restricted to western ...
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... predominance of I2a1b-M423 is surprising because it is completely absent elsewhere in CA/BA Europe, where the Y-DNA landscape is heavily dominated by R1b-M269 (Figs. 2- (Figs. 3 and 4). Even in Britain and Ireland, outside of Orkney most Neolithic Y-DNA lineages belong to haplogroup I2a2-S33 or I2a2a-M223 (Fig. 3), although, curiously, our Neolithic individual from Skye belongs to the very rare I2a2b-S154, seen elsewhere only in Middle Neolithic France (43). I2a1b-M423 seems to be largely restricted to western Neolithic Britain and Ireland, where it occurs rarely alongside I2a2a-M223, as well as I2a1a-CTS595 (41), which has not yet been found in ...
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... Europe after the end of the Neolithic. None are seen in post-Neolithic European archaeological remains. It is seen at only ∼1% in modern Britain and is almost absent in most of modern western Europe, although one recent subclade of I2a1b2-S392 has undergone dramatic expansion with Slavic populations in the Balkans (Figs. 2-4 and SI Appendix, Fig. S13) ...
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... and Orkney, most likely originating in Brittany (1, 49). I2a1b-M423 is seen in both Mesolithic and Neolithic Ireland, and the main cluster seen in Late Neolithic Ireland, I2a2a1a1-M284-found in the putative elite lineage at Newgrange-matches an Orcadian Neolithic lineage from the Isbister Chambered Cairn ("Tomb of the Eagles") on South Ronaldsay (Fig. 3 and SI Appendix, Fig. S13) (13). Both our data from BA Orkney and the Neolithic circumcoastal distribution of the Y-chromosome I2a1b-M423 haplogroup lend further support to this suggestion. European Neolithic society, at one extreme (but hardly peripheral) edge of its distribution, may have been patrilineal, patrilocal, and ...
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... originating in Brittany (1, 49). I2a1b-M423 is seen in both Mesolithic and Neolithic Ireland, and the main cluster seen in Late Neolithic Ireland, I2a2a1a1-M284-found in the putative elite lineage at Newgrange-matches an Orcadian Neolithic lineage from the Isbister Chambered Cairn ("Tomb of the Eagles") on South Ronaldsay (Fig. 3 and SI Appendix, Fig. S13) (13). Both our data from BA Orkney and the Neolithic circumcoastal distribution of the Y-chromosome I2a1b-M423 haplogroup lend further support to this suggestion. European Neolithic society, at one extreme (but hardly peripheral) edge of its distribution, may have been patrilineal, patrilocal, and hierarchical long before the arrival ...
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... We used the Pileup tool from GATK software (v.3.7.0) 69 to extract base calls over single-nucleotide polymorphism (SNP) sites in the 1,240k panel 75 for relevant genomes and selected one base call at random (base quality >30) for each site to generate pseudo-haploid genotypes. We merged 1,240k genotypes for 534 Iron Age individuals [17][18][19]41,61,[76][77][78][79][80] with a dataset of 5,326 modern individuals from western Europe 38,81 and, using approximately 266,000 sites common to both datasets, projected ancient genomes onto a PCA plot of modern variation using smartpca (version 16000) from EIGENSOFT 82 . We quantified EEF ancestry in British Iron Age genomes following a previously described procedure 17 . ...
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Roman writers found the relative empowerment of Celtic women remarkable¹. In southern Britain, the Late Iron Age Durotriges tribe often buried women with substantial grave goods². Here we analyse 57 ancient genomes from Durotrigian burial sites and find an extended kin group centred around a single maternal lineage, with unrelated (presumably inward migrating) burials being predominantly male. Such a matrilocal pattern is undescribed in European prehistory, but when we compare mitochondrial haplotype variation among European archaeological sites spanning six millennia, British Iron Age cemeteries stand out as having marked reductions in diversity driven by the presence of dominant matrilines. Patterns of haplotype sharing reveal that British Iron Age populations form fine-grained geographical clusters with southern links extending across the channel to the continent. Indeed, whereas most of Britain shows majority genomic continuity from the Early Bronze Age to the Iron Age, this is markedly reduced in a southern coastal core region with persistent cross-channel cultural exchange³. This southern core has evidence of population influx in the Middle Bronze Age but also during the Iron Age. This is asynchronous with the rest of the island and points towards a staged, geographically granular absorption of continental influence, possibly including the acquisition of Celtic languages.
... Conversely, a lack of admixture in Scotland is observed in the wake of the Middle to Late Bronze Age genetic turnover in Britain, suggesting that contact between Celtic speakers and now-unknown non-Celtic speakers could persist well into later prehistory (Patterson et al. 2022). In Orkney, Neolithic farmer lineages survived well into the Iron Age along the male line, and influx of Steppederived lineages mainly occurred along the female line (Dulias et al. 2022). ...
... A third possibility is to identify the Insular Celtic substrate with the few people descending from Neolithic farmers holding out in northern Britain and in Orkney (Dulias et al. 2022), but these holdouts seem rather peripheral to the more southerly location where Brittonic and Goidelic must have formed. ...
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We present a method for detecting evidence of natural selection in ancient DNA time-series data that leverages an opportunity not utilized in previous scans: testing for a consistent trend in allele frequency change over time. By applying this to 8433 West Eurasians who lived over the past 14000 years and 6510 contemporary people, we find an order of magnitude more genome-wide significant signals than previous studies: 347 independent loci with >99% probability of selection. Previous work showed that classic hard sweeps driving advantageous mutations to fixation have been rare over the broad span of human evolution, but in the last ten millennia, many hundreds of alleles have been affected by strong directional selection. Discoveries include an increase from ∼0% to ∼20% in 4000 years for the major risk factor for celiac disease at HLA-DQB1 ; a rise from ∼0% to ∼8% in 6000 years of blood type B; and fluctuating selection at the TYK2 tuberculosis risk allele rising from ∼2% to ∼9% from ∼5500 to ∼3000 years ago before dropping to ∼3%. We identify instances of coordinated selection on alleles affecting the same trait, with the polygenic score today predictive of body fat percentage decreasing by around a standard deviation over ten millennia, consistent with the “Thrifty Gene” hypothesis that a genetic predisposition to store energy during food scarcity became disadvantageous after farming. We also identify selection for combinations of alleles that are today associated with lighter skin color, lower risk for schizophrenia and bipolar disease, slower health decline, and increased measures related to cognitive performance (scores on intelligence tests, household income, and years of schooling). These traits are measured in modern industrialized societies, so what phenotypes were adaptive in the past is unclear. We estimate selection coefficients at 9.9 million variants, enabling study of how Darwinian forces couple to allelic effects and shape the genetic architecture of complex traits.
... Further aDNA studies have confirmed that the Late Iron Age Irish and northern British tribes, some of whom were later referred to as the Picts, were the direct descendants of these Beaker immigrants (Dulias et al., 2022;Margaryan et al., 2020). The Beaker Culture existed in Britain from 2500 BC until around 1800 BC (Bradley, 2007, p. 14), bracketing the constructions of Holme I and II. ...
Holme I and II were contemporary, adjacent Early Bronze Age (EBA) oak-timber enclosures exposed intertidally at Holme-next-the-sea, Norfolk, England, in 1998. Holme I enclosed a central upturned tree-stump, its function and intent unknown. Holme II is thought a mortuary structure. Both are proposed here best explained as independent ritual responses to reverse a period of severe climate deterioration recorded before 2049 BC when their timbers were felled. Holme I is thought erected on the summer-solstice, when the cuckoo traditionally stopped singing, departing to the ‘Otherworld’. It replicated the cuckoo’s supposed overwintering quarters: a tree-hole or the ‘bowers of the Otherworld’ represented by the tree-stump, remembered in folklore as ‘penning-the-cuckoo’ where a cuckoo is confined to keep singing and maintain summer. The cuckoo symbolised male-fertility being associated with several Indo-European goddesses of fertility that deified Venus - one previously identified in EBA Britain. Some mortal consorts of these goddesses appear to have been ritually sacrificed at Samhain. Holme II may be an enclosure for the body of one such ‘sacral king’. These hypotheses are considered, using abductive reasoning, as ‘inferences to the best explanations’ from the available evidence. They are supported with environmental data, astronomic and biological evidence, regional folklore, toponymy, and an ethnographic analogy with indigenous Late Iron Age practices that indirect evidence indicates were undertaken in EBA Britain. Cultural and religious continuity is supported by textual sources, the material record and ancient DNA (aDNA) studies.
... Bentley et al., 2002Bentley et al., , 2012Knipper et al., 2017), from aDNA analyses (cf. Schroeder et al., 2019;Dulias et al., 2022;Villalba-Mouco et al., 2022), or from a combination the two (Sjögren et al., 2020). At times these interpretations of female migration as evidence of patrilocality (residence) are linked to patrilineality (descent) and may even be seen as indications of patriarchy (power) (Mittnik et al., 2019;Sjögren et al., 2020). ...
Over the last several decades, the application of aDNA and strontium isotope analyses on archaeologically recovered human remains has provided new avenues for the investigation of mobility in past societies. Data on human mobility can be valuable in the reconstruction of prehistoric residential patterns and kinship systems, which are at the center of human social organization and vary across time and space. In this paper, we aim to contribute to our understanding of mobility, residence, and kinship patterns in late Prehistoric Iberia (c. 3300–1400BC) by providing new strontium data on 44 individuals from the site of Humanejos (Parla, Madrid). The study presented here is multi-proxy and looks at these new data by interweaving biological, chronological, and archaeological information. This analysis found that 7/44 individuals buried at Humanejos could be identified as non-local to the necropolis. Although more men (n = 5) than women (n = 2) were found in the non-local category, and more non-local individuals were identified in the pre-Bell Beaker (n = 5) than in Bell Beaker (n = 1) or Bronze Age (n = 1), we find no statistically significant differences concerning sex or time period. This contrasts with other archaeological datasets for late prehistoric Europe which suggest higher female mobility, female exogamy, and male-centered residential patterns were common. At Humanejos, we have also identified one non-local female whose exceptional Beaker grave goods suggest she was an individual of special status, leading to additional questions about the relationships between gender, mobility, and social position in this region and time period.
... Our understanding of more recent demographic changes in the British and Irish Isles-referring to Britain, Ireland and associated smaller islands-has also been expanded via large-scale sequencing of ancient genomes, revealing extensive gene flow from mainland Europe into southern Britain during the Middle Bronze Age, which contributed to genetic differentiation between Iron Age groups from southern and northern Britain [9,10]. Present-day genetic diversity in Wales, Cornwall, Devon and western Ireland indicates a long-standing genetic structure, possibly already present during the Iron Age [11], but the lack of ancient samples especially from Scotland limits our ability to directly test this hypothesis, and 'pockets' of older ancestries could have survived regionally in isolated populations for extended periods [12,13]. ...
... In terms of paternal Ychromosomal lineages, we assigned LUN004 to R1b-DF49 (Tables 1 and S6), which is predominantly distributed in the UK and Ireland and a sub-clade to R1b-P312/S116 haplogroup introduced to Britain by Bell Beaker peoples during the Chalcolithic, alongside steppe-related ancestry [7]. During the Chalcolithic, R1b-derived haplogroups largely replaced the predominant I2a Y-chromosome lineage in the British Neolithic [7,8,40] except in Orkney where I2a persisted into the Bronze Age [13]. R1b sub-clades are extremely common across Britain and western Europe from the Iron Age onwards [13,14,17,41]. ...
... During the Chalcolithic, R1b-derived haplogroups largely replaced the predominant I2a Y-chromosome lineage in the British Neolithic [7,8,40] except in Orkney where I2a persisted into the Bronze Age [13]. R1b sub-clades are extremely common across Britain and western Europe from the Iron Age onwards [13,14,17,41]. ...
There are longstanding questions about the origins and ancestry of the Picts of early medieval Scotland (ca. 300–900 CE), prompted in part by exotic medieval origin myths, their enigmatic symbols and inscriptions, and the meagre textual evidence. The Picts, first mentioned in the late 3rd century CE resisted the Romans and went on to form a powerful kingdom that ruled over a large territory in northern Britain. In the 9th and 10th centuries Gaelic language, culture and identity became dominant, transforming the Pictish realm into Alba, the precursor to the medieval kingdom of Scotland. To date, no comprehensive analysis of Pictish genomes has been published, and questions about their biological relationships to other cultural groups living in Britain remain unanswered. Here we present two high-quality Pictish genomes (2.4 and 16.5X coverage) from central and northern Scotland dated from the 5th-7th century which we impute and co-analyse with >8,300 previously published ancient and modern genomes. Using allele frequency and haplotype-based approaches, we can firmly place the genomes within the Iron Age gene pool in Britain and demonstrate regional biological affinity. We also demonstrate the presence of population structure within Pictish groups, with Orcadian Picts being genetically distinct from their mainland contemporaries. When investigating Identity-By-Descent (IBD) with present-day genomes, we observe broad affinities between the mainland Pictish genomes and the present-day people living in western Scotland, Wales, Northern Ireland and Northumbria, but less with the rest of England, the Orkney islands and eastern Scotland—where the political centres of Pictland were located. The pre-Viking Age Orcadian Picts evidence a high degree of IBD sharing across modern Scotland, Wales, Northern Ireland, and the Orkney islands, demonstrating substantial genetic continuity in Orkney for the last ~2,000 years. Analysis of mitochondrial DNA diversity at the Pictish cemetery of Lundin Links (n = 7) reveals absence of direct common female ancestors, with implications for broader social organisation. Overall, our study provides novel insights into the genetic affinities and population structure of the Picts and direct relationships between ancient and present-day groups of the UK.
... In general, molecular analysis of DNA could be a valuable method as TI is a common task in genetics [36]. However, DNA extraction from burnt bones is an expensive, destructive, time-consuming, and complex process that often does not lead to satisfactory results [37], especially as regards osteoarchaeological material [38]. ZooMS (Zooarchaeology by Mass Spectrometry) is affected by the denaturation and destruction of proteins caused by fire. ...
Differentiating cremated non-human bones from human ones in archaeological contexts is a challenging task. This analysis aims at proposing a rather solid criterion based on an osteoarchaeological sample. In this work, the main issues of taxonomic identification of cremated remains are analysed and a research methodology tested on an Italian protohistoric sample is proposed. The 314 subjects composing the sample come from 298 tombs of the Golasecca Civilization (1st millennium BC in north-eastern Italy). On a morphological basis, 246 bone fragments were selected from which as many thin sections were obtained for a histomorphological evaluation. From the analyses, we identified the presence of animals in burials, whereas a mere morphometric analysis was not able to recognize them. Furthermore, the taxonomic identification has allowed us to propose new hypotheses on the funerary rite of Golasecca linked to the zooarchaeological remains.
... According to the publications of Fóthi et al. [57], Neparáczki et al. [56], and Csányi et al. [16], the N1a1-M46, R1b-U106, and I2a-M170 lineages were the most widespread among the conquering Hungarians they examined. This suggests that the conquerors were of diverse origins, and while the N1a1-M46 subtype originated from the Ural region, the R1b-U106 (R1b1a1b1a1a1) lineage is known from the Late Copper Age/Early Bronze Age transition in Europe [79] and is most prevalent in Germanic-speaking people nowadays [80]. These observations fit the genomic data published by Maróti et al. [62], where haplogroups N1a1a1a1a2a1c (Y13850), N1a1a1a1a4 (M2128), I2a1a2b1a1a (YP189), and R1a1a1b2a (Z94) have been presented in notable frequencies. ...
Here we present 115 whole mitogenomes and 92 Y-chromosomal Short Tandem Repeat (STR) and Single Nucleotide Polymorphism (SNP) profiles from a Hungarian ethnic group, the Székelys (in Romanian: Secuii, in German: Sekler), living in southeast Transylvania (Romania). The Székelys can be traced back to the 12th century in the region, and numerous scientific theories exist as to their origin. We carefully selected sample providers that had local ancestors inhabiting small villages in the area of Odorheiu Secuiesc/Székelyudvarhely in Romania. The results of our research and the reported data signify a qualitative leap compared to previous studies since it presents the first complete mitochondrial DNA sequences and Y-chromosomal profiles of 23 STRs from the region. We evaluated the results with population genetic and phylogenetic methods in the context of the modern and ancient populations that are either geographically or historically related to the Székelys. Our results demonstrate a predominantly local uniparental make-up of the population that also indicates limited admixture with neighboring populations. Phylogenetic analyses confirmed the presumed eastern origin of certain maternal (A, C, D) and paternal (Q, R1a) lineages, and, in some cases, they could also be linked to ancient DNA data from the Migration Period (5th-9th centuries AD) and Hungarian Conquest Period (10th century AD) populations.
... Ancient DNA analysis was undertaken on 25 samples of human petrous bone and teeth at the University of Huddersfield Ancient DNA Facility. Twenty-three of these were successfully sequenced (Dulias et al. 2022) and yielded endogenous DNA content ranging between 3.1 and 66.5 per cent. Based on the genetic analysis, two infant samples from the multiple burial (which also had the same mitochondrial haplogroup and genetic sex) were identified as possibly being the same individual, and thus the minimum number of individuals analysed was 22. ...
... This suggests a substantial replacement of the Orcadian population between the Late Neolithic and the Bronze Age, probably as a result of migration from the British mainland at some point during the 'Beaker' period. There is, however, also clear evidence for continuity from the Neolithic in the male line of descent (Dulias et al. 2022), suggesting that patrilocal marriage patterns persisted from the end of the Neolithic into the Bronze Age. ...
... Genetic analysis provides us with a tantalising glimpse into possible Bronze Age inheritance and marriage systems. The persistence of indigenous male lineages for at least 1000 years after the end of the Neolithic (Dulias et al. 2022) may be associated with a system of patrilineal inheritance, in which household and land rights were passed down through the male line. Furthermore, that the number of households remained constant suggests that inherited 'assets' were not subdivided but remained intact through time. ...
The remarkable archaeological record of Neolithic Orkney has ensured that these islands play a prominent role in narratives of European late prehistory, yet knowledge of the subsequent Bronze Age is comparatively poor. The Bronze Age settlement and cemetery at the Links of Noltland, on the island of Westray, offers new evidence, including aDNA, that points to a substantial population replacement between the Late Neolithic and Bronze Age. Focusing on funerary practice, the authors argue for interconnecting identities centred on household and community, patrilocality and inheritance. The findings prompt are consideration of the Orcadian Bronze Age, with wider implications for population movement and the uptake of cultural innovations more widely across prehistoric north-western Europe.
Ancient microbial genomes can illuminate pathobiont evolution across millenia, with teeth providing a rich substrate. However, the characterization of prehistoric oral pathobiont diversity is limited. In Europe, only preagricultural genomes have been subject to phylogenetic analysis, with none compared to more recent archaeological periods. Here, we report well-preserved microbiomes from two 4,000-year-old teeth from an Irish limestone cave. These contained bacteria implicated in periodontitis, as well as Streptococcus mutans, the major cause of caries and rare in the ancient genomic record. Despite deriving from the same individual, these teeth produced divergent Tannerella forsythia genomes, indicating higher levels of strain diversity in prehistoric populations. We find evidence of microbiome dysbiosis, with a disproportionate quantity of S. mutans sequences relative to other oral streptococci. This high abundance allowed for metagenomic assembly, resulting in its first reported ancient genome. Phylogenetic analysis indicates major postmedieval population expansions for both species, highlighting the inordinate impact of recent dietary changes. In T. forsythia, this expansion is associated with the replacement of older lineages, possibly reflecting a genome-wide selective sweep. Accordingly, we see dramatic changes in T. forsythia's virulence repertoire across this period. S. mutans shows a contrasting pattern, with deeply divergent lineages persisting in modern populations. This may be due to its highly recombining nature, allowing for maintenance of diversity through selective episodes. Nonetheless, an explosion in recent coalescences and significantly shorter branch lengths separating bacteriocin-carrying strains indicate major changes in S. mutans demography and function coinciding with sugar popularization during the industrial period.
