Megalithic Tombs in Western and Northern Neolithic Europe Were Linked to a Kindred Society Supplement

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Megalithic Tombs in Western and Northern Neolithic Europe Were Linked to a Kindred Society Supplement Megalithic tombs in western and northern Neolithic Europe were linked to a kindred society Supplement Federico Sánchez-Quinto1*, Helena Malmström1,2*, Magdalena Fraser1,3*, Linus Girdland- Flink4, Emma M. Svensson1, Luciana G. Simões1, Robert George5,10, Nina Hollfelder1, Göran Burenhult3, Gordon Noble6, Kate Britton6,7, Sahra Talamo7, Neil Curtis6, Hana Brzobohata8, Radka Sumberova8, Anders Götherström9, Jan Storå10#, Mattias Jakobsson1,2# * Equally contributed to this work #corresponding authors Affiliations 1 Human Evolution, Department of Organismal Biology, Uppsala University, SE-752 36 Uppsala, Sweden 2Centre for Anthropological Research and Department of Anthropology and Development Studies, University of Johannesburg, P.O. Box 524, Auckland Park, 2006, South Africa 3Department of Archaeology and Ancient History, Uppsala University-Campus Gotland, SE-621 67 Visby, Sweden 4Research Centre in Evolutionary Anthropology and Paleoecology, School of Natural Sciences and Psychology, Liverpool John Moores University, Byrom Street, Liverpool, L3 3AF, UK 5Royal Prince Alfred Hospital, Sydney Australia 6Museums and Special, Collections, University of Aberdeen, Sir Duncan Rice Library, Bedford Road, Aberdeen, AB24 3AA, Scotland 7Department of Human Evolution, Max Planck Institute for Evolutionary Anthropology, Deutscher Platz 6, 04103, Leipzig, Germany 8Institute of Archaeology of Czech Academy of Sciences, Letenská 4, CZ-11801 Prague, Czech Republic 9Archaeological Research Laboratory, Department of Archaeology and Classical Studies, Stockholm University, SE-106 91 Stockholm, Sweden 10Osteoarchaeological Research Laboratory, Department of Archaeology and Classical Studies, Stockholm University, SE-106 91 Stockholm, Sweden Data Availability Raw sequencing reads produced for this study have been deposited in the European Nucleotide Archive (ENA) under accession number ENA: PRJEB31045. www.pnas.org/cgi/doi/10.1073/pnas.1818037116 Supplement: Megalithic tombs in western and northern Neolithic Europe Contents S1 Archaeological Background ............................................................................................................... 3 S2 The ancient individuals, provenance and site description .................................................................. 5 Carrowmore, County Sligo, Ireland .................................................................................................... 6 Primrose Grange, County Sligo, Ireland ............................................................................................. 9 Midhowe, Orkney, Scotland.............................................................................................................. 13 Lairo, Orkney, Scotland .................................................................................................................... 14 Balintore, Ross and Cromarty, Scotland ........................................................................................... 14 Ansarve, Gotland, Sweden ................................................................................................................ 15 Kolin, Bohemia, Czech Republic ...................................................................................................... 18 S3 Sample preparation, DNA extraction and Library preparation ........................................................ 20 S4 Processing of NGS data ................................................................................................................... 22 S5 Contamination estimation ................................................................................................................ 24 S6 Calling mtDNA haplogroups ........................................................................................................... 26 S7 Calling Y chromosome haplogroup ................................................................................................. 28 S8 Molecular sexing of individuals ...................................................................................................... 36 S9 Reference panels and group labels ................................................................................................... 38 S10 Estimating kinship relationships among Megalith individuals ...................................................... 40 S11 Population genetic analyses ........................................................................................................... 55 S12. Primrose Grange, Tomb. Original field documentation. ............................................................... 70 References supplement ..................................................................................................................... 75 2 Supplement: Megalithic tombs in western and northern Neolithic Europe S1 Archaeological Background On farmer migrations routes along the Atlantic facade Recent archaeogenetic research has shown the importance of human migrations and mobility in the dispersal of Neolithic lifeways over the European continent. Apart from the Early Neolithic inland and the Mediterranean coastal expansion routes, archaeological research suggests a western European Atlantic maritime route connecting Neolithic cultures via the Atlantic, the North Sea, and the Baltic littoral (1–4), of which some routes may have been important already during the Mesolithic period as well as the Early Neolithic (5, 6). Genetic patterns are generally consistent with archaeological data (7–14), although the processes of population expansion and migration resulted in some degree of genetic differentiation between different Neolithic groups (15). While available genetic data from Neolithic farmers that were buried on the northwestern fringes of Europe (i.e. the British Isles and Scandinavia) show strong affinities to Central European farmers (16–21), individuals buried in Megalithic chambered tombs in Ireland (Ballynahatty), and Sweden (Gökhem) had strong affinity to Neolithic and Chalcolithic Iberian farming groups (19, 22). However, the extent to which coastal and maritime migration routes existed along the islands and the coastal regions of western Europe remains unknown. Megalithic burial tradition Megalithic tombs (from the Greek megas, great and lithos, stone) are large burial constructions usually built from rock, soil and wood (23). Based on both materials and design, different types of megalithic tombs can be identified, e.g. dolmens, passage graves, and court cairns to name a few. All types may collectively be categorized as chambered tombs and/or megalithic tombs. These types of burials are often found along the former coastlines and on islands, suggesting maritime connections (24–26); however, inland locations also occur. The geographic and chronological distributions of the different types of chambered tombs vary. While dolmens and passage graves are commonly found across the whole Atlantic façade and some Mediterranean islands (24–30), chambered cairns are mainly found in the British Isles (31, 32). However, the explanation behind the compelling similarities in the construction and design of dolmens and passage graves from Iberia to southern Scandinavia, including the British Isles, remains elusive. The inter-regional interaction linking the west European megalithic sites, consistent also with the dispersal of domesticated resources, raw materials and arte- facts, has often been attributed to shared social and cultural systems, and also cosmology (23, 31, 33). The European monumental burials, which accompanied funerary rites, have been studied at least since early medieval times (23). The stone structures are problematic to date, and many burials do not contain organic remains that can be directly radiocarbon dated. However, the earliest dates seem to fall in the middle of the 5th millennium BCE in Brittany and the Channel Islands, in Catalonia, in 3 Supplement: Megalithic tombs in western and northern Neolithic Europe west and central southern France, on Sardinia, in northern Italy, and possibly in Apulia and also on Corsica (33). An important insight to the complexity regarding the dating of various phases in megalithic tombs in Ireland is a recent study and re-examination of the find materials from a multi- phase passage tomb on Baltinglass Hill, Co. Wicklow, excavated in the 1930s (34). The study is important and presents evidence of unexpectedly early megalithic construction, possibly as early as 3900 Cal BCE, challenging conventional views of the emergence of the Irish Neolithic. At least three phases between c. 3900-2900 Cal BCE can be determined at Baltinglass Hill (34). Also, a comprehensive study of all available dates from the Mesolithic-Neolithic interface, the Neolithic and the Bronze Age in Ireland was recently published, showing that traces of Neolithic activities are scarce prior to 3750 Cal BC (35). Megalithic tombs are, as a rule, multiperiod sites. They have been built, used, rebuilt and reused again. These burials were often collective graves, and their usage also continued for centuries and/ or millennia, including secondary burials in later time periods (21, 24, 26, 36–41). During additional deposits, earlier material and human remains have often been pushed aside or probably even thrown out from the chambers. Consequently, the dating of the primary construction is most often difficult but very important when it comes to understanding the underlying settlement-subsistence system and economy of the tomb-building society in question. There are
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