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Major migration events in Holocene Eurasia have been characterized genetically at broad regional scales1–4. However, insights into the population dynamics in the contact zones are hampered by a lack of ancient genomic data sampled at high spatiotemporal resolution5–7. Here, to address this, we analysed shotgun-sequenced genomes from 100 skeletons spanning 7,300 years of the Mesolithic period, Neolithic period and Early Bronze Age in Denmark and integrated these with proxies for diet (13C and 15N content), mobility (87Sr/86Sr ratio) and vegetation cover (pollen). We observe that Danish Mesolithic individuals of the Maglemose, Kongemose and Ertebølle cultures form a distinct genetic cluster related to other Western European hunter-gatherers. Despite shifts in material culture they displayed genetic homogeneity from around 10,500 to 5,900 calibrated years before present, when Neolithic farmers with Anatolian-derived ancestry arrived. Although the Neolithic transition was delayed by more than a millennium relative to Central Europe, it was very abrupt and resulted in a population turnover with limited genetic contribution from local hunter-gatherers. The succeeding Neolithic population, associated with the Funnel Beaker culture, persisted for only about 1,000 years before immigrants with eastern Steppe-derived ancestry arrived. This second and equally rapid population replacement gave rise to the Single Grave culture with an ancestry profile more similar to present-day Danes. In our multiproxy dataset, these major demographic events are manifested as parallel shifts in genotype, phenotype, diet and land use.

Western Eurasia witnessed several large-scale human migrations during the Holocene1,2,3,4,5. Here, to investigate the cross-continental effects of these migrations, we shotgun-sequenced 317 genomes—mainly from the Mesolithic and Neolithic periods—from across northern and western Eurasia. These were imputed alongside published data to obtain diploid genotypes from more than 1,600 ancient humans. Our analyses revealed a ‘great divide’ genomic boundary extending from the Black Sea to the Baltic. Mesolithic hunter-gatherers were highly genetically differentiated east and west of this zone, and the effect of the neolithization was equally disparate. Large-scale ancestry shifts occurred in the west as farming was introduced, including near-total replacement of hunter-gatherers in many areas, whereas no substantial ancestry shifts happened east of the zone during the same period. Similarly, relatedness decreased in the west from the Neolithic transition onwards, whereas, east of the Urals, relatedness remained high until around 4,000 BP, consistent with the persistence of localized groups of hunter-gatherers. The boundary dissolved when Yamnaya-related ancestry spread across western Eurasia around 5,000 BP, resulting in a second major turnover that reached most parts of Europe within a 1,000-year span. The genetic origin and fate of the Yamnaya have remained elusive, but we show that hunter-gatherers from the Middle Don region contributed ancestry to them. Yamnaya groups later admixed with individuals associated with the Globular Amphora culture before expanding into Europe. Similar turnovers occurred in western Siberia, where we report new genomic data from a ‘Neolithic steppe’ cline spanning the Siberian forest steppe to Lake Baikal. These prehistoric migrations had profound and lasting effects on the genetic diversity of Eurasian populations.

The transitions from foraging to farming and later to pastoralism in Stone Age Eurasia (c. 11-3 thousand years before present, BP) represent some of the most dramatic lifestyle changes in human evolution. We sequenced 317 genomes of primarily Mesolithic and Neolithic individuals from across Eurasia combined with radiocarbon dates, stable isotope data, and pollen records. Genome imputation and co-analysis with previously published shotgun sequencing data resulted in >1600 complete ancient genome sequences offering fine-grained resolution into the Stone Age populations. We observe that: 1) Hunter-gatherer groups were more genetically diverse than previously known, and deeply divergent between western and eastern Eurasia. 2) We identify hitherto genetically undescribed hunter-gatherers from the Middle Don region that contributed ancestry to the later Yamnaya steppe pastoralists; 3) The genetic impact of the Neolithic transition was highly distinct, east and west of a boundary zone extending from the Black Sea to the Baltic. Large-scale shifts in genetic ancestry occurred to the west of this “Great Divide”, including an almost complete replacement of hunter-gatherers in Denmark, while no substantial ancestry shifts took place during the same period to the east. This difference is also reflected in genetic relatedness within the populations, decreasing substantially in the west but not in the east where it remained high until c. 4,000 BP; 4) The second major genetic transformation around 5,000 BP happened at a much faster pace with Steppe-related ancestry reaching most parts of Europe within 1,000-years. Local Neolithic farmers admixed with incoming pastoralists in eastern, western, and southern Europe whereas Scandinavia experienced another near-complete population replacement. Similar dramatic turnover-patterns are evident in western Siberia; 5) Extensive regional differences in the ancestry components involved in these early events remain visible to this day, e

Reindeer hunters on the moveSmall Lateglacial sites in eastern JutlandThe Søvind site is an example of a Late­glacial site discovered during the excavation of much later remains. Given its limited size and depth beneath the soil surface, it is a type of locality that is unlikely ever to have been found by “usual” archaeological methods.Only scattered Lateglacial finds have been recorded from the central part of eastern Jutland to date (fig. 22). These probably do not provide a true picture of the settlement of the time, but reflect chance situations. The two new sites presented here, Søvind and Elhøj, are therefore very significant.The Søvind site is situated on a “land bridge” between high ground extending south towards Horsens Fjord, a large east-west oriented valley in Lateglacial times, and higher hills to the north (fig. 1). There are no watercourses or lakes in the vicinity, and its location sets the site apart from the majority of known Lateglacial sites, which are frequently situated by fresh water.Stratigraphy. Beneath the plough soil was a c. 40-50 cm layer of sand, which became increasingly brownish-yellow and reddish-brown with depth (fig. 2). The flint artefacts were found within this sand layer and in two pits (fig. 3). There were no secondary glacial changes, for example solifluction etc., of the archaeological deposits. The only cultural remains were of flint. The find-bearing deposits were sieved through a fine sieve and delimited in all directions by find-free areas (fig. 3). The flint was found in particular in two pits in the middle of the flint scatter (fig. 3), and outside these the artefact concentration was only c. 1-2 pieces/m2. The flint covered an area of c. 9.5 x 4 m (c. 38 m2) (fig. 3). The area of the Søvind site is less than that of other Lateglacial sites, for example Bro I in northwest Funen, which covered an area of c. 50 m2, but corresponds to that of the Lateglacial site of Langå I.Settlement features. In the middle of the flint s

Analyser af dyreknogler fra en submarin boplads, Ronæs Skov, ved Gamborg Fjord på Fyn, påviser mindst syv arter af fisk, fire arter af fugle og nitten arter af pattedyr. Fiskeriet har især været domineret af torsk. Fuglefangsten har bestået af skarv, svane, spurvehøg og natugle. Foruden den for perioden karakterisktiske jagt på vildsvin, kronhjort og rådyr har der foregået en vis hvalfangst. Det er interessant, at der er fanget marsvin, som der også i nutiden har været drevet storjagt på i dette område, Analyses of animal bones excavated on a submarine stone Age settlement, Ronæs Skov, at Gamborg Fjord, Funen. At least seven species of fish, four species of birds and nineteen species of mammals were exploited at this Ertebølle site. Fishing for cod were characteristic and swans as well as cormorants were hunted. The most frequent mammals at the site in the Ronæs skov material are wild boar, red deer and roe deer. A significant element of marine mammals is also apparant: several species of seal and whale. Especially bones from harboar porpoise are worth noting.

A flaker from the Dogger Bank in the North Sea By the end of the Ice Age, the present North Sea was dry land stretching from Denmark to the British Isles. Here, life conditions for hunter tribes must have been good and similar to those in the adjoining areas – Southern Scandinavia, Eastern England, and the Netherlands. Due to the rise of the world seas, this large land area was gradually flooded after the Ice Age, and recent geological investigations have made it possible to gain a relatively good picture of the development history of the North Sea during the last c.14000 years. These investigations show that the highest land areas – primarily the Brown Bank towards the south and the Dogger Bank towards the north – must have been large land areas far into the early Stone Age. As opposed to our geological knowledge, the archaeological finds fail to provide information about the settlement structure of the early Stone Age in this area. However, they are important because they can provide information about important issues such as when and to which extent the different land areas were flooded by the sea, and the nature of the cultural affiliations of the hunter groups living there.So far, the Stone Age finds have almost all been made in the southern part of the North Sea (Brown Bank), whereas there are just a few finds from the rest of the large sea territory. A few of these artefacts are connected to the Maglemose Culture, whereas the rest can either not be dated or are subject to the discussion whether they are actually artefacts or just products of nature.However, a new find now helps throwing light on some of these questions. It is a pressure flaker fished out of the sea at Dogger Bank, at a depth of 30 to 40 metres. The tool was made from a tine of red deer (Cervus elaphus) antler, the point of which was truncated at an angle and shows clear marks of pressure and wear (Fig. 1). In North Europe, this type of artefact is known only from the late Maglemose Culture se

Activity sites from the Ertebølle periodDyngby III and Sindholt Nord In order to understand the maritime culture of the Ertebølle period in Denmark (5600-4000 BC), it is necessary to take as many sources as possible into account. In the following, two new aspects of coastal Ertebølle finds, represented by the Dyngby III and Sindholt Nord sites, are described (Figs. 1 and 10). Dyngby III is located at the foot of a hill bordering an area which in the Stone Age was a small bay connected with Kattegat (Fig. 1). Several small sites have been excavated along this prehistoric coastline – both “Køkkenmøddinger” (shell middens), and coastal sites without any shell deposits; to these can now be added a new type of coastal site, which is the central issue of this article. The site is rather small (10 m x 10 m), and the culture layers were encapsulated by tufa deposited by fresh water. Charcoal powder, flint debris, and shells from oysters and cockles define the culture layers. No settlement structures were recorded, but the presence of charcoal, burnt flint, and burnt bones indicate the use of fire on the beach.The find material is very restricted. Apart from worked flint, it comprises just a few potsherds, “pot-boilers”, and small bone splinters. The number of artefacts and the range of types represented are limited (Figs. 4, 5, and 6). The most common artefact types are triangular or discoid flint pieces with irregular scars along the edges (Fig. 7) – types which are rare on other Ertebølle sites. The heavy wear indicates that they were used as hammers on a hard material such as bone or antler, but as there are no artefacts or waste in the find material to support these observations, the explanation for the use and occurrence of these tools at Dyngby III is open to discussion. In addition, the numerous core renovation flakes and blades demonstrate an extensive production of blades on this site. The absence or near-absence of blade-tools (scrapers and borers, as well as a ve

A magnificent ornamented axe from the Early Ertebølle Period Even in Denmark, Mesolithic antler axes with geometric patterns covering the whole surface are a rarity. The ornamented tools from the Early Ertebølle Period in Western Denmark have been published previously (S. H. Andersen 1981), and this article should be considered a supplement to this past work. The axe shown in fig. 1 was excavated from a sub-marine Culture layer at Skærbæk in Eastern Jutland, fig. 7. Consequently, it is a settlement find. The raw material is red deer antler. The axe had been scraped completely smooth and then given a geometric ornamentation covering the whole surface. The main part of the ornamentation had been scratched in a fine and thin line, but a small part had been carried out as a wider and coarser line. The coarser ornamentation is secondary, as it covers the fine line. Ornamentation carried out in two phases has been recorded previously on several other antler axes from Jutland from the Ertebølle Period. The dominant motif is a regular chess­board pattern, which is repeated in different variations, fig. 1. Analysis of the ornamentation shows -as is the case for several other Danish antler axes- that it was not carried out at the same time but over a considerable period. On the axe from Skærbæk at least 15 different "pattern episodes" can be identified, which partly overlap and therefore must represent a quite long "pattern making period", fig. 2. This theory is supported by the fact that the axe is very polished with use and tremendous grinding. The net motif is a characteristic ornament of the Early Ertebølle Period, but it is usually carried out in a rhomboid pattern. The regular chessboard ornament on the Skærbæk axe is rare. The other motifs are known from other Jutland antler axes. According to type, ornamentation and find circumstances, the Skærbæk axe can be dated to the Early Ertebølle Period, c. 5400- 4700 BC. This date matches other finds of ornamented Ertebølle ax,                           &nbsp

Ertebølle harpoons and killer whale teeth Aspects of marine hunting in the Ertebølle period Since the Danish Ertebølle harpoons were presented together nearly twenty-five years ago (Andersen 1972, 1976 and 1983), figs. 1-8, 12 and 19-20, several new finds have been made, a number of which are from excavations of well-dated settlement finds, figs. 9-11 and 13-18. Moreover, harpoons have now been found in Scania, northern Germany and in northern Poland, so they now cover the entire Ertebølle cultural province around the western Baltic, figs. 22-23. Most of the finds are, as previously, from coastal settlements and the sea, but several new finds also come from inland tracts. Further harpoons derive from areas from which several finds were already known, for example from the Little Belt area and Horsens Fjord, figs. 22-23. It is also interesting that an increasingly large number of harpoons made of whale bone have been found, and waste from whale-bone harpoon production is also known from several new settlement finds, figs. 25 and 28. Among the new finds there is also a patterned harpoon of type C from the settlement excavation at Lystrup Enge (early Ertebølle culture), fig. 16. Corresponding ornamented harpoons were previously known only from Horsens Fjord; a harpoon of this kind has subsequently been found at Podjuch near the mouth of the Oder, fig. 19. The original division into 4 main types, types A, B, C and D, has been further strengthened with the latest finds, figs. 22-23. In addition, it is now possible to define a small group comprising an even older type, A-1, represented by the harpoons from Tudeå and Skateholm, fig. 15; this group must be dated to the latest Kongemose culture or the transition between the Kongemose and Ertebølle cultures, c. 5500/5400 BC. Among the new finds it is also remarkable that a harpoon has now for the first time been found in a grave from Skateholm II in Scania, fig. 15. This find really documents the importance of the harpoon at t,                               &nbsp

Ertebølle canoes from Lystrup In Late Atlantic times (6000-4000 B.C.), the time of the Ertebølle Culture, the coast near Århus was much more sinuous than it is today. About 8 km north of Århus there was a 5.5 km long and 1.5 km wide inlet, now damp meadows and cultivated fields called Lystrup Enge (fig. 1). This inlet seems to have come into existence at around 6000-5500 B.C., and many coastal settlements, particularly those dating from the older part of the Ertebølle Culture have been recorded along its shores. One of the discoveries is bones of a killer whale (Orcinus orca) (fig. 12). One of the settlements, which was excavated after 1981 but is not yet published, is the largest known Jutland settlement of the older Ertebølle culture (fig. 2), and the most remarkable of its finds are the canoes described as Lystrup 1 and Lystrup 2 (S.H. Andersen 1994). The site lies on the southern side of a low gravel point running east-west parallel with the coast. The huts stood on the point itself, where only flint now survives, but the rubbish dumps lay in nearby waterlogged fjord deposits, where animal bones, antler, and much wood is also present, the latter consisting especially of many long hazel stakes, which were the remains of fishing fences (fig. 4). The water beside the settlement was not only dump, but also a fishery with food waste, making it one of the oldest kitchen middens in the country. Not only the location, but also the many fishbones and mollusc shells show the importance of the sea for life at the site. So do the bones of small and large whales. The commonest fish were cod, flatfish (plaice, dab, etc.) and eels. Fishing was done close inshore using permanent fixed gear, primarily in the form of traps. The presence of bones of mackerel, garfish, and eels show chat some at lease of the fishing took place in the summer and early autumn. The whales species present were bottle-nosed dolphin, dolphin, white­beaked dolphin, and killer whale, as well as the larger,                    &nbsp

APPENDIX A Revised Dating of Layers 3 and 4 from the Norslund Settlement In Kuml 1965 there was a presentation of the Norslund settlement (1). This included the publication of two C-14 dates of layer 3, which gave an average dating to 3755 ± 120 BC in C-14 years (K-990 and K-991). This article also discussed the great similarity in the archaeological material between layer 3 and the layer below it, layer 4, which was C-14 dated to 4470 ± 130 BC in C-14 years (K-993). In 1965, however, the latter dating was called into question due to the fact that the dated material might have consisted of redeposited older material. Since 1965 new excavations of the Ertebølle sites on Jutland and Funen and typological studies of the artefact content of C-14 dated cultural layers such as Brovst (2), Haldrup Strand, Ø. Jølby Kær, Skellerup Vig, Meilgård, and Tybrind Vig (3) have all shown the published C-14 datings of layer 3 to be incorrect. In contrast, the dating of Norslund layer 4 does correspond well with later results. The C-14 tests from Norslund did not derive from the cultural layers in the main area of the excavation but from a marine layer series in an isolated trial trench below the settlement (4). With the aid of small profiles, an attempt was made to connect profile D of the main area with profile H of the trial trench, although direct contact between the profiles was not achieved. The lower part of layer 4, however, has definitely been found in the profiles and the layer is identical with profile H, layer 2, from which the dating K-993 K-993 + 4470 ± 130 BC derives. In 1965, Norslund layer 3 was identified with profile H layer 6, which was dated by means of two tests of charcoal (hazel) to 3780 ± 120 and 3730 ± 120 BC (K-990 and K-991). However, layer 6 was only seen in profile H, and the layer completely tapered out in the northern end of the profile toward the main area. Thus there cannot have been any direct contact with layer 3, and consequently there is no indisp,                          &nbsp

&nbsp, Patterned Oar Blades from Tybrind Vig During excavation of the submerged Ertebølle settlement Tybrind Vig (4200-3300 BC) on the Little Belt (fig. 1 and 2) (1), a number of characteristic wooden implements were recovered, of a kind not previously found in Denmark, fig. 3. Similar objects are known from the Rüde 2 settlement at Satrup, South Schleswig, where they are called "spades" (2). In this article, however, these implements are considered to be oars. They are made of ash and have a c. 100-120 cm long shaft with a broad heart-shaped blade, fig. 3. During the 1980 excavation, the patterned oar was recovered, and another one was found later, fig. 7. Patterned wooden objects have never before been found in a Danish Mesolithic settlement and the new find reveals and entirely new art form and technique. The oar fig. 4 is made of ash (Fraxinus excelsior) and lacked on excavation only a little of the blade. The shaft is 116 cm long and the blade 20 cm wide (original width c. 30-35 cm), 30 cm long an about 1 cm thick. The surface is smooth and exhibits no traces of working or wear. One well preserved side of the blade is patterned. The pattern covers the top two-thirds of the blade an appears in low negative relief in the smooth surface. When the oar was found, it was inlaid with a brown pigment, fig. 4. Careful investigations suggest that the pattern has been pressed into the surface of the wood, for instance by means of a stamp or the like. The point of inlaying a dark pigment in the pattern must have been to emphasize its effect, a technique which is known from patterned Ertebølle objects in other materials, for instance antler, bone and amber (3). The composition covers the width of the blade and is symmetrical about the long axis and a transverse axis, fig. 4a-b. The ornament consists of a mixture of geometrical motifs and "softer", sinuous bands, balancing the effect of strict composition and the geometrical motifs. The pattern is built around a transverse central

&nbsp, Ertebølle Art: New Finds of Patterned Ertebølle Artefacts from East Jutland In view of the abundance of finds from the many Ertebølle settlements which have yielded detailed insight into various important aspects of this phase of the Stone Age, it is surprising that patterned objects appear so rarely. Moreover, as the scanty finds -with few exceptions- either derive from earlier excavations or are stray finds, it has not been possible to date them precisely in relation to the culture. This infrequency of patterning has therefore been considered a characteristic feature of the Ertebølle culture and has often been mentioned to exemplify the difference between this period and the earlier Maglemose and Kongemose cultures. Today this conception can no longer be said to have general validity. One of the areas which most clearly indicates this is eastern Jutland, where, despite the many Ertebølle finds with favourable conditions of preservation, only very few patterned artefacts have been published. During the past ten or fifteen years, however, investigations of Ertebølle sites in eastern Jutland have indicated that the ornamented objects are more common than earlier assumed. At the same time, the new finds also show an interesting variation with regard to selection of raw materials, motives, composition, and patterning technique. Finds have been made of artefacts of deer antler, bone, amber, pottery, and stone. In addition, the find contexts of several of the objects have made it possible for the objects -and with them the patterns and the pattern techniques- to be dated more precisely within the period. At the present, however, we cannot determine with certainty whether the new finds simply reflect a special feature of the Ertebølle settlements of this region or whether the entire find material hitherto made in Jutland has not been representative. Perhaps the answer lies in a combination of these possibilities. This article offers an overview of the newly found objects

New harpoon finds After the publication in KUML 1971 of the harpoons then known (1) my attention was drawn to a number of other specimens, hitherto unknown and all to be found in private collections.Notable among the new harpoons is the single-rowed specimen fig. 1, which is a casual find from Tudeå, western Zealand. Found in fresh water, it was probably used for catching large freshwater fish like salmon, pike or catfish.Apart from superficial recent damage, the harpoon is entire. The raw material is red deer antler. The surface has been scraped smooth and the base exhibits a row of distinct secondary cuts which are probably due to a change in the method of hafting during the life of the harpoon. The line hole has a biconical section. The overall length is 32.6 cm. The two decorative elements seen on the convex, smooth side in this specimen - fine lines and deeper nicks - are combined and therefore contemporaneous. The patterns consist of lines with short transverse strokes (Clark motif c), bands with transverse hatching (Clark motif f) with small triangular figures which occur here for the first time in the Danish Mesolithic, and large hatched triangles.The decoration is difficult to interpret on account of its condition; it is probably purely geometric, but the possibility that it is a stylized animal or human representation cannot be ruled out. The ornaments on the Tudeå harpoon are, with one exception, well known from previous finds. Corresponding motifs are known from both the Maglemose and Kongemose cultures, especially from the many axes of the latter, for example those from Ølby Lyng (Kemisk Værk) (25), Carstensminde (26) and Værebro Å (28). The Tudeå harpoon should on the basis of its similarity to other well-dated specimens be assigned to the later Kongemose - earliest Ertebølle culture.The disharmony between the patterns and the shape of the harpoon shows that the decoration was probably not made for the harpoon. As several instances are known of broken

Aggersund. An Ertebølle settlement on the Limfjord The Ertebølle culture in Denmark is probably one of the best studied mesolithic cultures in northern Europe. Archaeological investigations have, however, concentrated almost exclusively on the large settlements, particularly their middens, while excavations of the smaller settlements of the culture have only rarely been undertaken or published. Large sites, representing lengthy occupation, are not suited to the elucidation of conditions in the short term. Small settlements on the other hand open the possibility of erecting a finer chronological system, which can be used in the analysis of the larger sites, as well as the possibility of studying more short term, seasonal subsistence activities. Finally the small sites make it possible to check the degree to which subsistence affects the tool kit present. This situation can be shown by means of an example from Aggersund in north Jutland. The area round Aggersund is now some 6 m higher in relation to the sea than it was during the stone age (2), during which period it was broken up into a series of larger or smaller islands separated by narrow straits. or broader shallower stretches of water, fig. 1. Traces of the action of the stone age sea are much in evidence in the form of old seacut cliffs, raised beach systems and areas of raised Littorina seabed (3). Because of the particular topographic and geological situation Aggersund was the narrowest point of the fjord in the prehistoric period as well. West and northwest of Aggersund are areas of fen -which in the stone age were wide areas of shallows- where large quantities of waterbirds collect while migrating through in the spring and autumn. In this part of the Limfjord it should therefore be possible to find settlements which exploited these waterbird migrations to some considerable degree. The excavation of a small Ertebølle settlement on the north side of the fjord at the farm »Bundgård« near Aggersund in the centr,             &nbsp