In 3100 BC, hundreds of years before the first Indo-Europeans would set foot in Scandinavia, two very different groups of people had been maintaining highly contrasting ways of life for at least 400 years. Despite often living in very close proximity to each other. Everything ranging from their spiritualism, to their burials and even diets differed greatly. As did their DNA.

The maritime-oriented hunter-gatherers of the Pitted Ware culture (PWC), descendants of the old foragers behind the Kongemose, Slate Culture and Ertebølle traditions held a tight grip of the coastline all the way from Southwestern Norway to the Åland islands in modern day Finland, to Northern Jutland and Northeastern Zealand in Denmark. They also inhabited Eastern Svealand in Sweden, the islands of Gotland and Öland, and the Western coast of Sweden along with the zone around the large Lake Vänern in Western Sweden. The people of the Pitted Ware culture had a strong preference for coastal areas, but evidence of significant inland settlement also exists. Some of these areas were not particularly well suited for the needs of the other group, the agricultural and pastoralist Funnelbeaker people, while others were sought after by people of both cultures.

Currently, only very low levels of admixture between contemporaneous Scandinavian Hunter-Gatherers (SHG) from the Pitted Ware culture and Early European Farmers (EEF) from the Funnelbeaker culture (FBC) have been noted. This article will go through the variation in farmer-hunter relations and what kind of genetic implications they might have had.

One archaeological site that is dramatically different from other contemporaneous sites in Scandinavia is the Alvastra pile dwelling.

The pile dwelling at Dagsmosse

Out in rural Östergötland in Sweden lies the spring mire of Dagsmosse in which the remains of this well-preserved pile dwelling are located. The dwelling is situated at the foot of a mountain by the name of Omberg (known in the past as Ammobiærh, literal translation being “Misty Mountain”). The slopes of the mountain face the large Lake Vättern to the west. This mountain has a rich mythological past, and in the past it was often associated with elves, giants and trolls, all commonly featured in Norse mythology. In local folklore, many tales about the mountain and the area around it exists. Nobel laureate Verner von Heidenstam wrote a great collection and summary of folk tales from the region back in 1914.

One of the more commonly mentioned tales is that of the Geatish queen Omma, who is said to have lived on Omberg. The queen had many admirers, one of whom was the red haired giant known as Ringabergsbusen. This giant lived on the other side of Lake Vättern, and was desperate to propose to Omma. One day, in winter when the ice was frozen solid, he decided to give it a shot, and mounted his massive horse. He rode across the ice at full speed on his great steed, but was unable to rein his mount in, and collided with the mountain. The giant and his horse were pulled down to their deaths into the depths of Lake Vättern, never to emerge again. In local folklore, Ringabergsbusen’s collision with the mountain is said to have left its mark on the landscape, causing the distinct cracks in the mountain that are visible to this day. Queen Omma’s tears are said to cause the mist that often surrounds the mountain. Mythology aside, this mountain and the area surrounding it is of vast archaeological importance. The mountain has several hill forts dating to the Migration Period and the largest of them is situated at the very top of the mountain. One of the hill forts is even named after old Queen Omma. Below is a figure showcasing the area.

It was in the spring mire near the foot of this mystical mountain that the Alvastra pile dwelling was built in 3100 BC. During the time of what in Scandinavian archaeology is called the Early Middle Neolithic, when the dwelling was at its peak, logs were piled all over the ground and formed a floor of sorts that rose above the ground of the mire. In the middle of the mire a 600 m2 large square platform made out of logs was placed. The platform was supported by more than a thousand piles that had been pushed down into the mire. They seem to have been placed there to prevent people on the outside from seeing what went on inside rather than to serve as a defensive palissade, as indicated by their sparse positioning. When the Neolithic inhabitants of Scandinavia built defensive palissades, they placed the piles much more densely and deeper into the ground than they did at Alvastra. A causeway of approximately 100 metres length served as the only entrance and exit to the pile dwelling. The dwelling itself had two main sections, both of which contained 8 or 9 separate rooms. The rooms had plenty of hearths made out of limestone and clay.

Back in those times, raising such a structure was far from easy work. Not only did it require a whole lot of work being put in to it, it also required a lot of people. Archaeologist Hans Browall (Browall 1986) estimated that the construction would have required a large community of farmers, likely around 20 households and hundreds of people in order to be completed. Using flint and stone axes, the Stone Age people there cut and piled the logs to create the pile dwelling. At the time the pile-dwelling was constructed, the farmers of the Funnbelbeaker culture had already been forced to give up a lot of ground to the expanding Pitted Ware foraging culture. Some of the farmers migrated south again, taking the route through Denmark and possibly ending up back in Germany, which had been the main starting point of their initial expansion to Scandinavia. Even so, some of them persisted in parts of southern Scandinavia (especially in the inland), despite the hunter-gatherers of the Pitted Ware culture being on the warpath. One group of FBC people that had remained in Scandinavia was most likely behind the construction of the pile-dwelling at Alvastra, judging by the style of construction and the tools required to make a structure like it.

Speaking of Funnelbeaker culture tools, plenty of those have been found at Alvastra’s pile dwelling. The pottery recovered at the site, however, is mainly of the Pitted Ware type. This is astonishing, because this is arguably the only site in Scandinavia where such conclusive evidence of an intimate cohabitation between these two cultures can be found. Most of the pottery and tools found at Alvastra had not been produced at the site, but had been carried there from nearby settlements. If we get back to the previously mentioned limestone hearths, we can find even more proof of this cohabitation. Residue from sustenance typical of the Funnelbeaker culture, such as barley and wheat, nuts and apples, are found. As are bones from the cattle that was vital for the FBC lifestyle. In addition to these rather expected finds, however, residue of sustenance typical of the Pitted Ware culture is also found around the hearths. Fish like perch and pike, as well as bones from bigger game that was most likely brought down by PWC foragers, such as bear, wolf and moose.

The red deer and pig bones at the site could easily be attributed to either culture, because while the FBC people did hunt deer on occasion they are not known to have hunted bear or moose, unlike the PWC. Pitted Ware people did keep pigs, although it was a highly supplemental practice and nowhere near as much of a staple for them as it was for the Funnelbeaker people. It is worth noting that Pitted Ware people in Denmark more often kept pigs than the Pitted Ware people in Sweden and Norway, likely due to more close interactions with farmers. Their Ertebølle culture predecessors (roughly 5800-4000 BC) had practiced animal husbandry to some degree as well, even though they were mainly hunter-gatherers and rather thriving ones, at that.

Looking at the discoveries at Alvastra, one can only really make one conclusion. The pottery and tools combined with the varied dietary remains strongly indicates that these 2 groups resided at Alvastra at the same time.

The pile dwelling itself could have been used for anything, and archaeologists have speculated endlessly about its actual purpose. Some popular suggestions are, for example, that it was used as a place to keep food for cattle during the winters, or that it was a center for trade where various groups of hunter-gatherers and farmers could meet on “neutral” ground and exchange goods. The most popular theory among Scandinavian archaeologists, however, has always been that the Alvastra pile-dwelling site was a religious center, and a highly important one. Uppsala is believed to have been one of the main religious centres of the Norse. Perhaps Alvastra had a similar function as the temple that would stand at Uppsala in the Viking Age. Evidence suggests that the pile dwelling was mainly inhabited during the summer and early autumn season, which could give some credence to the theory that the pile dwelling at Alvastra was connected to a seasonal religious ritual of some kind. It is possible that these festivities were related to either the harvest season for the farmers, or to one of the more prestigeous hunting seasons for the hunters. A frequently encountered theory in Scandinavian archaeology is that human sacrifice was performed at Alvastra. There is no solid proof of this, but considering that the piles that supported the platform were not placed for defensive purposes, but rather to prevent insight, it seems reasonable to assume that the frequenters did not want people outside to see what was going on on the inside. Whether or not that was because they were performing human sacrifice inside is left for anyone’s imagination.

Despite Alvastra being the only pile dwelling in Sweden and despite its apparent status as an important meeting place during the Early Middle Neolithic, the site was not in use for very long. Evidence suggests that during the final period of use, the Alvastra pile dwelling was seemingly no longer used by the people of the Funnelbeaker culture. Instead, it was mainly used for burials by the Pitted Ware foragers, who praticed single or double grave inhumation with the dead lying on their back, often with red ochre placed inside the burial. They seem to mostly have buried grown men and boys at Alvastra, for some reason. Perhaps the site had a great religious importance for them, possibly having something to do with manhood or rites of passage related to it.

One young male was found to have been scalped, and his skull was found on a stake just outside of the pile dwelling. Scandinavian Hunter-Gatherers were sometimes known to put the heads of dead people on stakes. The most well known example of this are the skulls at Kanaljorden, Motala where the skulls mounted on the stakes also happen to be quite well preserved. Below is an image of what it could look like when the Motala SHGs were in such a mood:

This act could have served as a way to intimidate rival groups, but it has been speculated that it could also have been done as a religious act to fend off evil spirits. Naturally, we will never know exactly why they did it, but archaeology makes it clear that during the final period prior to the abandonment of the Alvastra pile dwelling, Pitted Ware people seem to have been its sole inhabitants. This of course, raises the question if the Funnelbeaker people at some point lost access to Alvastra’s sacred grounds, or if they willingly stopped coming there. At a quick glance, the former seems like the likelier theory here, as this happened roughly around the same time that the PWC people were undertaking an expansion and replacement of the FBC that seems surprisingly organized and has some striking similarities to the resurgence of Western-Hunter-Gatherers in Middle Neolithic Western Europe. The Funnelbeaker people were pushed south, and so was Europe’s northernmost border of agriculture. However, Browall 1986 stresses that agriculture was still practiced, albeit on a much smaller scale than before, in the same region at the time of Alvastra’s abandonment, and after it. It’s quite reasonable to assume that pockets of FBC people endured the PWC expansion in this area and simply lost their dominance in the region and perhaps as a result of that, also lost their access to the pile dwelling at Alvastra.

Archaeogenetic research on the skeletal remains from the pile dwelling is currently underway, and should be able to give us even more information about the relationship between the farmers and the foragers at Alvastra. Of course, Alvastra is not the only place where the two cultures interlapped, it is simply the only site where they seem to have done so relatively harmoniously, at least in the beginning.

A segregated island: Funnelbeaker farmers and Pitted Ware hunter-gatherers on Gotland

On the island of Gotland in the Baltic Sea these two populations coexisted in a rather peculiar fashion. They lived very close to each other on a relatively small island but still did not interact or mix with each other much. Fraser et al. (2018) emphasized the rather consistent differences in dietary patterns between the FBC and the PWC. Much like on the mainland, the people of the PWC on Gotland almost exclusively got their food from maritime sources, whereas the people of the FBC had a much more land-based diet and only in some cases seem to have eaten maritime foods. Perhaps the hunter-gatherers were not fond of the FBC farmers venturing too close to their fishing waters.

Pitted Ware individuals on Gotland had low levels of EEF admixture which they got from the FBC. However, FBC people on Gotland for the most part do not seem to have had any SHG admixture from the PWC. This may suggest that the little mixing that did occur between these two cultures happened in such a way that a small number of individuals from the FBC were incorporated into the PWC. Whether this may have occurred through the kidnapping of wives from the FBC, or from FBC people voluntarily taking up a hunter-gatherer lifestyle is hard to tell. Evidence of violent conflict between forager groups and agricultural groups on Gotland during this period exists, just as it does on the mainland during the same period. The PWC males on Gotland all appear to have carried I2 haplogroups, more specifically of the I2a1 branch. Typical SHG lineages, in other words, and ultimately WHG-derived. Interestingly, Funnelbeaker males from a megalith grave at Ansarve on Gotland were also found to carry I2a1 haplogroups. A Funnelbeaker sample (Gökhem4 from Skoglund et al. 2014) from Västergötland in Sweden also belonged to haplogroup I2a1. However, the FBC men likely got their Y-DNA somewhere between their migration from Germany into Scandinavia, not directly from local SHGs like those of the Pitted Ware culture.

Sanchéz-Quinto et al. (2019) showed that FBC farmers at both the Gökhem and Ansarve sites in Sweden showed signs of hunter-gatherer admixture on the autosomes as well as the X chromosome, which indicates that they had recent admixture from a HG source that was not necessarily strictly male-biased. This suggests that the (slight) admixture that took place between these groups of FBC and PWC happened on somewhat even terms. At least that’s what the current, albeit limited, data suggests. Had they gotten their I2a1 lineages from SHGs, higher amounts of SHG admixture would have accompanied said haplogroups even after a couple of generations, but the FBC samples lack the necessary EHG ancestry that would come with SHG ancestry (even though they do have trace amounts of EHG ancestry, with an outlier having as much as ~6%) and as such, it seems more likely that the FBC people got their WHG-rich admixture from a population further to the south. The farmers from Ansarve on Gotland have trace amounts of EHG admixture, while 2 Funnelbeaker samples from Rössberga on the mainland have either noise levels of EHG (<0.6%) or 0. The latter is also true for the FBC samples from Gökhem.

Likewise, Pitted Ware samples from Gotland also carried small amounts of EEF admixture, but not enough to suggest that there was any continous admixture between them and the FBC people on the island.

Considering the duration during which these two populations lived closely to each other on Gotland, their interactions and mingling seem to have been remarkably limited. We can guess endlessly about what it was that what made them so reluctant to mingle, but judging by archaeological findings from Scandinavia as a whole, the Pitted Ware people and the Funnelbeaker people on Gotland and Öland do not seem to have liked one another. Plenty of evidence of violence between the two groups exists, they deliberately lived in different areas even when inhabiting the same (limited in size, in the case of Gotland) regions, and barely any admixture took place between the groups. This raises the question if PWC-FBC relations were this poor everywhere in Scandinavia, or if there were exceptions. It is an especially interesting question in regards to genetics. Was there a SHG-EEF hybrid population similar to the WHG-Anatolian Neolithic Farmer populations found in Western, Central and Southern Europe?

Djursland, Denmark

Sites from Djursland are useful because they enable us to understand what it looked like when the Pitted Ware culture was on the move, how they became dominant in areas previously belonging to the Funnelbeaker culture, and how organized they were. At the Kainsbakke site, near Kolindsund, PWC people were found to have reused old TRB ritual sites. At Kainsbakke, PWC cylindrical blade cores and tanged arrowheads of all 3 PWC types have been found in and around FBC-type megalith graves. This could mean that the PWC met the FBC on peaceful terms at this site, but it could also mean that the FBC were simply driven away and that for some reason, the PWC foragers liked to reuse their old ritual sites. However, no defensive palissades built by the FBC have been found at the site.

At Kainsbakke, both animal bones and artefacts were imported from other PWC settlements, mainly western Sweden, but also Gotland and inland parts of Jutland. The area surrounding the site had large deposits of high-quality flint, which is believed to have been acquired and imported to PWC communities with a lack of access to flint deposits of comparable quality. Iversen et al. (2016) stresses that even though flint was a highly sought after resource in both the FBC and the PWC, it was probably sought after for different reasons. The hunter-gatherers would have wanted long flint blades in order to produce top notch hunting equipment, or tools for warfare. For the FBC these long flint blades would primarily have been used for harvesting activities. The pattern of settlement at Kainsbakke shows that it served as a more permanent settlement from which provisions were taken to smaller, more seasonal camps. Pitted Ware activity in a region tended to have this pattern, often being based out of a permanent settlement from which expeditions were sent out and seasonal camps set up, often with signficant geographic distance from the core settlement.

In northeastern Djursland, what appears to be another place of religious importance for the Funnelbeaker people can be observed, but it has few similarities to Alvastra. According to Klassen et al. (2020) the 6 kilometer long boggy valley was likely a sacred place for the Funnelbeaker people inhabiting the region. Two streams flow in the valley, and they are believed to have deposited sacrificial items in both. What is so interesting about this sacred valley is that despite it being located in between two of the main clusters of PWC sites in northeastern Djursland, the PWC foragers seem to have completely avoided the valley. While the valley, with its swamps and bogs, has ideal conditions for hunting, the hunters simply appear to have steered clear of it. Klassen et al. attributes this to the fact that they might have been aware of the valley’s ritual importance to the FBC. This is peculiar, because we could be looking at a case of Pitted Ware people being unusually respectful of the boundaries set by the FBC and of their customs. While the lack of PWC wetland deposits and signs of PWC settlement in the valley could indicate that the relations between the PWC and FBC were better in this area, it could of course also simply be a coincidence. However, if we assume that there was a mutual respect and exchange, then there could also have been an exchange of genes and ideas.

Differences between Ertebølle and Pitted Ware in reaction to agriculture, and a brief look at the Norwegian Pitted Ware sphere

The Ertebølle culture is of interest when discussing interactions between the Funnelbeaker culture and the hunter-gatherers of Scandinavia, as the Ertebølle hunter-fishers interacted a great deal with the FBC. The main difference between Ertebølle and Pitted Ware in this regard is that the Ertebølle people were quite willing to adopt agriculture in many cases, and seem to have mingled a lot with the FBC. The Pitted Ware people, however, for the most part appear to have been quite reluctant to switch to a sedentary way of life, much preferring the hunt and journeying the waters which they had thoroughly mastered. At a quick glance, it’s easy to think that the Pitted Ware people were simply backwards and unwilling to adapt to new technology. The reality is that they were highly adaptable but also incredibly selective. They took what they likely perceived as being the only good aspects of the Funnelbeaker culture, like animal husbandry, and in some cases such as in Åland (Vanhanen et al. 2019) the PWC even cultivated crops, although to a limited extent.

Was there really a Neolithic in Norway?

– Christopher Prescott, Professor of Archaeology at the Universtity of Oslo

In Norway, the Pitted Ware hunter-gatherers are believed to have had knowledge of the Neolithic lifestyle due to contacts with the FBC people who may have been living in the Oslofjord region, but they still preferred to keep their earlier (mostly) Mesolithic lifestyle. The evidence of agriculture associated with the Funnelbeaker culture in Norway is very sparse. This may in part be due to the acidic nature of the soil in the regions of Eastern Norway where the FBC is believed to have existed, as it often destroys traces of old cultivation. Even so, the distribution of Funnelbeaker type axes in this region does hint at a farmer presence, although in theory it could just as likely have been the result of the hunter-gatherers in the region either obtaining such axes through trade, or adopting FBC technology.

The hunter-gatherers in this region were indeed very picky, and chose to adopt some Neolithic technology associated with the production of ceramics and in some cases flint axes, but seem to have dismissed the rest of the Neolithic package. They basically appear to have lacked any kind of interest in switching to an agricultural or pastoralist mode of production. SHG DNA persisted for a long time in northern Norway, as proven by sample VK531 from Troms (Margaryan et al. 2020), a man who had a Mesolithic genetic profile despite living around 2400 BC. This was likely the case in northern Sweden as well. However, these remnant forager populations, at least the ones in the far north, did not make significant genetic contributions to later Scandinavians.

Agriculture in Norway was arguably much more successfully introduced during its second wave, brought there permanently by the agro-pastoralist Late Neolithic migrations. Interestingly, it is also in the Late Neolithic that Norway experienced the increase in population size that agriculture brought on much earlier in places where early agriculture took off to a greater extent.

Another significant difference between the Ertebølle culture foragers and those of the Pitted Ware culture is that the Ertebølle people were far more limited in mobility (as stated in Andersen 2018 and Klassen et al. 2020). The PWC was very mobile and undertook expeditions that stretched across much larger distances than the movements of the Ertebølle people ever did. Ertebølle people were also significantly fewer in numbers and their territory was far more geographically limited than the PWC’s, and their settlements were also less centralised. Generally speaking, Pitted Ware hunter-gatherers were more organized.

In terms of genetics, only 1 Ertebølle DNA sample exists, and it’s a somewhat dubious one due to being recovered through questionable methods (oral microbiome from chewed birch pitch.) However, what the genome from Jensen et al. (2019) tells us is that this particular Ertebølle girl was far more WHG-shifted than Pitted Ware people were, as the latter had their own genetic profile which they had inherited from their ancestors, known in archaeogenetics as SHG which was a mix of WHG and EHG. The female sample from Jensen et al was an unmixed WHG. It is possible that the EHG component was not present in Denmark to a significant degree before the Pitted Ware culture expanded there. However, 1 sample is not enough to conclude anything about the genetic profile of Ertebølle foragers. It is quite likely that Ertebølle in Scania had more EHG admixture, and it is also possible that EHG admixture was present in Danish Ertebølle as well. The Funnelbeaker people in northern Europe got their HG-admixture from an unmixed source of WHG, which could definitely have been Ertebølle in the case of the FBC in Scandinavia. This WHG-rich farmer ancestry is present in significant amounts in modern Scandinavians as well.

The Western part of the FBC horizon in parts of the Netherlands and Germany may have been even more rich in WHG, having the bulk of their WHG ancestry derived from remnants of the Swifterbant culture foragers. But that’s a different story.

As mentioned above, very few Ertebølle remains have been sampled for DNA, but we can look at farmers in Germany who most likely had direct admixture from Ertebølle foragers in Denmark. Namely, farmers from the Middle Neolithic Walternienburg-Bernburg culture. This culture is believed by some archaeologists to have sprung out of influence from Denmark, and this was recently supported by a DNA study. The authors of Beau et al. (2017) suggested that this culture had its origins in migrations out of Scandinavia. The Walternienburg-Bernburg culture was rich in hunter-gatherer mtDNA haplogroups, similar to some FBC groups in Scandinavia. Additionally, Haak et al. (2015) analyzed the Y-DNA haplogroup of one male from the Walternienburg-Bernburg culture and found that he carried I2a1, just like the Funnelbeaker males in Scandinavia did. If the people of the Walternienburg-Bernburg culture actually were migrants (or back-migrants, rather) from Scandinavia, it is likely that they received their WHG-rich admixture from Ertebølle foragers, and possibly their I2a1 Y-DNA haplogroups as well. This gives us at least some idea of what kind of paternal lineages the Ertebølle foragers could have carried, Ahrensburg-derived WHG lineages.

The willingness of the Ertebølle foragers to be incorporated into the expanding Funnelbeaker culture means that later Ertebølle samples will most likely have very significant EEF admixture. Even so, the Ertebølle people were very few in numbers and it is therefore plausible that the bulk of the WHG admixture in Scandinavian Funnelbeaker people came from admixture that they acquired before entering Denmark and the rest of Scandinavia.

It is not known what triggered the seemingly organized expansion of PWC. It could have been a growth in hunter-gatherer population size, but it could also have been a result of growing tensions between the PWC and the FBC, perhaps due to religious or other cultural differences. Another option is that the Pitted Ware hunter-gatherers were forced south by another group of more northern foragers infringing on their territory. In practice, what the PWC expansion meant was that a fairly rapid “de-Neolithisation” took place as hunter-gatherers moved in to areas that were previously only inhabited by the FBC. Violence was often involved in this expansion, and FBC settlements were abandoned. The subsistence shifts from agro-pastoralism to hunting, often with a focus on sealhunting. These shifts are usually accompanied by the appearance of Pitted Ware-style pottery.

When it comes to adaptability, Pitted Ware foragers in some places like on Gotland did not have any trouble with adopting new customs such as burial traditions from later populations in Scandinavia, such as the Indo-European Battle Axe culture. Malmström et al. (2020) showed that Pitted Ware foragers with no steppe admixture were buried in Battle Axe-influenced graves, despite no geneflow between the two groups at the time. The question is why the PWC people were willing to imitate Battle Axe culture traditions but unwilling to do the same with Funnelbeaker traditions. It is possible that since the PWC were able to not only compete with the FBC, but even outcompete them and replace them in places despite having a much lower population size, they did not see the Funnelbeaker cultural package as a desirable way of life. The Battle Axe culture also had a much larger population size than the PWC did, but were far more dominant in the region than the FBC ever managed to become. Perhaps the more warlike and expansionistic nature of the Battle Axe culture was perceived by the PWC as more desirable to imitate than the FBC. The PWC did end up being completely assimilated into the Battle Axe culture, and their relatively small population either disappeared or left only small traces in later Scandinavians. Despite this, some Pitted Ware customs related to seafaring and to hunting and warfare (arrowheads) persisted for a long time, well into the Scandinavian Late Neolithic and even into the Nordic Bronze Age.

Plague: a factor to consider for Forager-Farmer relations?

Something that should not be neglected when discussing close interactions and cohabitation between the PWC foragers and the FBC farmers is disease. Before 2018, it was often believed that it was the migrations of Yamnaya-related peoples from the steppe that introduced the plague in Europe. This turned out to be incorrect. Rascovan et al. (2018) discovered that a FBC woman found at Gökhem carried a strain of the plague-inducing bacterium Yersinia pestis. The authors of the study proposed that the FBC farmers would have picked up this disease from contacts with/origins in the larger EEF settlements in central and southeastern Europe. While it is certainly possible that Yersinia pestis was present (at a significant amount, even) in the large farmer settlements in continental Europe, the most recent data does not necessarily suggest that this was something that was unique to farmer settlements, or even originated in them. Susat et al. (2021) found Y.pestis in a 5000-year old hunter-gatherer from Latvia. Now, despite Y. pestis clearly being present even among hunter-gatherers in the eastern part of the Baltic that had no direct contact with large EEF populations, it is more likely that it would be present at a higher rate in more densely populated regions inhabited by farmers. Therefore it is also quite fair to infer that it was the Funnelbeaker people who were the main carriers of Y. pestis in Scandinavia at the time of the Alvastra pile dwelling.

The PWC foragers on Gotland and Öland did engage in trade with foragers in Latvia and Lithuania, but Y. pestis has not been found among them so far. The PWC on the mainland did not have direct contact with foragers in the Baltic states, and would thus have been less likely to contract the plague on their own. Because of that, a presence of Y.pestis among FBC people in and around Alvastra could have meant a serious harm for the PWC people and if contracted and spread, been detrimental to Pitted Ware demographics, given their low population size and likely poor resistance against that type of disease. This is of course something to consider even when discussing hunter-farmer interactions outside of shared sites like Alvastra. Did the hunters know about the plague, and avoid farmer settlements because of that? It is something to ponder.

What the pile-dwelling at Alvastra could mean for archaeogenetics

It is clear that the nature of the interactions between foragers and farmers in Scandinavia varied enormously depending on the location. In most places, relations appear to have been rather poor and both violence and segregation/isolation between groups were commonplace. Some exceptions exist, with the Alvastra pile dwelling being the best example, and these exceptions will be very important for getting the full picture of what actually happened. Pitted Ware foragers were, for the most part, not particularly eager to take on the full Neolithic package. In many places, like southern and western Norway for example, they had a thriving community of coastal fishing settlements and most likely felt no need to switch to a fully Neolithic mode of production. The same thing applies to most of the Pitted Ware culture in Sweden.

A team of researchers at Uppsala University in Sweden are currently carrying out a multidisciplinary study of the skeletons from the Alvastra pile dwelling. This study will examine not only DNA, but also go into greater detail about the health, diet, kinship and population size about the people that resided at the Alvastra pile dwelling. The ancient DNA results will likely turn out to be quite illuminating in regards to what exactly happened during the last phase of Alvastra’s Neolithic inhabitance, and if the cohabitation between the PWC and the FBC there also meant that there was more admixture between them than in other places, where admixture between them seem to have been either very slight or non-existent. Another benefit of genetic data from Alvastra will be that for the first time in the aDNA record, we will likely get Y-DNA haplogroups from Pitted Ware hunters from mainland Scandinavia. By closely examining dietary patterns, researchers will be able to see if a hypothetical admixture between Pitted Ware and Funnelbeaker culture individuals resulted in a more mixed dietary pattern, i.e. genetically HG-shifted individuals subsisting on a grain-based diet, or EEF-shifted individuals with hunter-gatherer diets.

The archaeological findings from East-Central Sweden and from Åland hint at a slightly less one-sided relationship between the PWC and the FBC than in, for example, Scania in southern Sweden or on Gotland and Öland. In the former, the FBC simply disappeared for the most part, while the FBC in the latter two places had very limited interactions with the PWC despite often living in their proximity. In East-Central Sweden there was significant cohabitation and some exchange of ideas and technology but the archaeological record also reveals that ultimately, de-Neolithisation occurred and many Funnelbeaker settlements disappeared.

Given the evidence of close exchange between FBC and PWC in some places, it is likely that admixture will also have taken place. Finding answers to if this potential admixture resulted in a new population that positioned itself on a cline between SHG and EEF, or if it only happened occassionally and only resulted in a few genetic outliers will be of great importance for further understanding of the population structure in Late Neolithic Scandinavia. By looking at Y-DNA, researchers will be able to determine if the PWC expansion was similar to the WHG resurgence in Middle Neolithic Europe where a very male-biased resurgence of WHG ancestry took place. Will the upcoming study of DNA at Alvastra reveal a male-biased hunter-gatherer admixture, as was often the case in Middle Neolithic Europe? Or will the results show a more balanced admixture? There are many gaps that need to be filled in when it comes to population dynamics during this time period.

The upcoming research will also be examining the presence of Yersinia pestis as well as other disease status among the Alvastra dwellers. This will also be able to clear things up in regards to what it was that led to the pile dwelling at Alvastra eventually being abandoned. Was it disease, violence between groups, both, or neither?

Getting to the bottom of PWC:FBC admixture, continuity and relations is also vital to comprehending their interactions with the later Corded Ware-derived Battle Axe culture that would enter Scandinavia only a couple of hundred years after the abandonment of the Alvastra pile dwelling. We already have quite a bit of data from PWC contemporary with the BAC on Gotland, but what exactly was the Battle Axe culture up against in mainland Scandinavia? Time will tell.

Naturally, this article will be updated as soon as the results from the upcoming study by Malmström and colleagues are out.

References:

Nielsen et al., (2019): De-Neolithisation in southern Norway inferred from statistical modelling of radiocarbon dates https://www.sciencedirect.com/science/article/abs/pii/S0278416518300825?dgcid=author

Prescott (1999): Was there really a Neolithic in Norway?https://www.cambridge.org/core/journals/antiquity/article/abs/was-there-really-a-neolithic-in-norway/905A2611AEF092918871916AE25A668F

Iversen, Rune(2016) : The Transformation of Neolithic Societie.s. An Eastern Danish Perspective on the 3rd Millennium BC https://www.tandfonline.com/doi/abs/10.1080/00293652.2016.1230145

Susat et al., (2021): A 5,000-year-old hunter-gatherer already plagued by Yersinia pestis https://www.cell.com/cell-reports/fulltext/S2211-1247(21)00645-8

Rivollat et al., (2017): Multi-scale ancient DNA analyses confirm the western origin of Michelsberg farmers and document probable practices of human sacrifice https://journals.plos.org/plosone/article?id=10.1371/journal.pone.0179742

Brandt et al., (2013): Ancient DNA reveals key stages in the formation of Central European mitochondrial genetic diversity https://www.ncbi.nlm.nih.gov/pmc/articles/PMC4039305/

Fraser et al., (2018): New insights on cultural dualism and population structure in the Middle Neolithic Funnel Beaker culture on the island of Gotland https://www.sciencedirect.com/science/article/abs/pii/S2352409X17303231?via%3Dihub

Gummesson et al., Cambridge University Press (2018): Keep your head high: skulls on stakes and cranial trauma in Mesolithic Sweden https://www.cambridge.org/core/journals/antiquity/article/keep-your-head-high-skulls-on-stakes-and-cranial-trauma-in-mesolithic-sweden/39BD3070DCC745A9CBAB1DF52FEC2782

Wilhelmson et al., (2009): An analysis of the bone material from the Alvastra megalithic tomb https://portal.research.lu.se/portal/sv/publications/an-analysis-of-the-bone-material-from-the-alvastra-megalithic-tomb(98e5e4b7-c59c-4739-bac2-e5f28b3f810e).html

The Dolmen in Alvastra, by Janzon, Gunborg O, Torbjörn Ahlström, Mattias Ahlbeck, Göran Skog, Helene Wilhelmson, Alan Crozier https://www.biblio.com/book/dolmen-alvastra-janzon-gunborg/d/936533005

Klassen et al., (2020): Pitted Ware Culture Settlement on Djursland https://www.academia.edu/42795202/Pitted_Ware_Culture_Settlement_on_Djursland?email_work_card=title

Sánchez-Quinto et al (2019): Megalithic tombs in western and northern Neolithic Europe were linked to a kindred society https://www.ncbi.nlm.nih.gov/pmc/articles/PMC6511028/

Jensen et al., (2019): A 5700 year-old human genome and oral microbiome from chewed birch pitch https://www.nature.com/articles/s41467-019-13549-9

Rascovan et al., (2018): Emergence and Spread of Basal Lineages of Yersinia pestis during the Neolithic Decline https://www.cell.com/cell/fulltext/S0092-8674(18)31464-8?_returnURL=https%3A%2F%2Flinkinghub.elsevier.com%2Fretrieve%2Fpii%2FS0092867418314648%3Fshowall%3Dtrue

Einar Østmo (2007): The Northern periphery of the TRB graves and ritual deposits in Norway https://onlinelibrary.wiley.com/doi/abs/10.1111/j.1600-0390.2007.00102.x

Skoglund et al., (2014): Genomic Diversity and Admixture Differs for Stone-Age Scandinavian Foragers and Farmers https://science.sciencemag.org/content/344/6185/747

Vanhanen et al., (2019): Maritime Hunter-Gatherers Adopt Cultivation at the Farming Extreme of Northern Europe 5000 Years Ago https://www.nature.com/articles/s41598-019-41293-z

Elin Andersson (2018): The first passage tombs in Falbygden https://www.diva-portal.org/smash/get/diva2:1217754/FULLTEXT01.pdf

Hans Browall (1986): The Alvastra pile dwelling; a social and economic base http://libris.kb.se/bib/7608412

Malmström et al., (2020): The Neolithic Pitted Ware culture foragers were culturally but not genetically influenced by the Battle Axe culture herders https://onlinelibrary.wiley.com/doi/10.1002/ajpa.24079

Sørensen et al., (2012): The expansion of agrarian societies towards the North – New evidence for agriculture during the Mesolithic/Neolithic transition in Southern Scandinavia https://www.researchgate.net/publication/236874462_’The_expansion_of_agrarian_societies_towards_the_North_-_New_evidence_for_agriculture_during_the_MesolithicNeolithic_transition_in_Southern_Scandinavia

Malmström et al., (2009): Ancient DNA Reveals Lack of Continuity between Neolithic Hunter-Gatherers and Contemporary Scandinavians https://www.cell.com/current-biology/fulltext/S0960-9822(09)01694-7

© Genomic Atlas 2021