Europe's Astonishing Hunter-Gatherer Resurgence

By 6,500 years ago, Neolithic farmers had colonised almost every part of Europe. From the Aegean to the Atlantic coast, people grew domesticated crops like wheat and peas, and raised livestock like sheep, pigs and cattle. The ancient hunter-gatherers of Europe had been pushed to the fringes and were in danger of disappearing altogether. But then after about 4500 BC, everything changed. It's common to think of Europe's prehistory as a straight line, from hunter-gatherers to farmers and from stone tools to bronze and iron. But recent genetic research has revealed that this story is actually more complicated, and that some of the people who first called Europe home, the hunter-gatherers who had lived here since the end of the Paleolithic, did not simply vanish when agriculture arrived. In fact, thousands of years after the first farmers spread across the continent from the Near East, traces of Europe's older inhabitants re-emerged in the DNA of later communities. And it wasn't just an increase in hunter-gatherer ancestry. Many of the Neolithic farmer male lineages were also replaced with hunter-gatherer ones. This unexpected genetic signal has been called the late Neolithic hunter-gatherer resurgence. So how and why did this happen? For me, this is one of the most intriguing stories in all European prehistory, and it perhaps challenges the popular narrative of complete population turnovers and the total lack of continuity between the Mesolithic populations of Europe and those that came after. This is the inspiring story of Europe's hunter-gatherer resurgence. It's only thanks to the science of ancestral DNA analysis that the late hunter-gatherer resurgence has been revealed to us. And that's why on this video, I've partnered with MyHeritage. For a limited time, you can now get your own DNA kit from MyHeritage for the incredibly low price of $25 euros or pounds, depending on where you live, using my link and code. I've never seen an ancestry DNA testing kit for this low of a price before, so if this is something you're interested in doing, then you should grab this now using my link while it's this price. You might remember that I previously partnered with MyHeritage and shared the results of my ethnicity estimate. The new algorithm for the ethnicity estimate is now updated to version 2.5. I'm a British man with an Irish grandparent, and my ethnicity estimate reflects that. And I've added new ethnicities and reference panels, which gives you more detail than ever. But to me, the most exciting new feature they've introduced is called Ancient Origins. This lets you analyze your own DNA by comparing it to specific ancient reference populations from various eras. Looking at this Ancient Origins breakdown for the Bronze Age era, divided between these two reference populations, I am 59% Neolithic farmer and 41% Western stepherder, and you can get breakdowns for different eras. The most relevant one for this video subject divides my ancestry by ancient West Eurasian reference populations. This is a really exciting opportunity to engage with our ancient past in a unique and individualized way. With my code DAN25, you're not only getting this ridiculous price on the DNA kit, when you order it, you're also getting a 30-day free trial, free shipping, and the Ancient Origins feature. And much more I don't have the time to tell you about. So to find out your own Ancient Origins, click the link in the description box or scan the QR code on screen and use the coupon code DAN25 to get your DNA testing kit for the insanely low price of $25 pounds or euros. Thank you very much to MyHeritage for sponsoring the video. At the end of the Upper Paleolithic, as glaciers covered the north, Europe's hunter-gatherers retreated to refugia in the south, such as in Iberia and the Balkans. Around 11,700 years ago, the glaciers began to retreat, and those hunter-gatherer populations recolonized the continent from those refugia. And quickly, these various communities adapted to the post-glacial landscape. Archaeologists call them Mesolithic societies, Mesolithic being the Middle Stone Age, coming between the Paleolithic Old Stone Age and the Neolithic New Stone Age. The Mesolithic hunter-fisher foragers adapted to a wide range of environments across the continent. They fished in rivers and along coasts, hunted red deer and wild boar, and gathered nuts, especially hazelnuts, and berries in the forests. They also needed other resources like antlers, and especially stone like flint, with which to make their tools. Geneticists have now tested hundreds of ancient individuals and classified them as different populations. Most commonly, these have been called the Western Hunter-Gatherer, or WHG population, a distinct genetic cluster that extended from Iberia to the Baltic. And in Eastern Europe, they defined a different population with different ancestry, they called Eastern Hunter-Gatherer, or EHG. These populations mixed with each other across the regions they interacted, so these two populations actually blended together to one degree or another, forming a cline in the middle. In Scandinavia, this mixed population has been given its own designation, Scandinavian Hunter-Gatherer, or SHG. But then, after about 7000 BC, farming in Europe began in the Aegean, after spreading from Anatolia. It then spread across Europe in two main waves, one across the Mediterranean coasts all the way to Iberia, the other through the Balkans and into Central Europe. This brought a new way of life, where people grew domesticated crops and raised livestock. Archaeologists saw this wave of new pottery and tool types, houses and settlements, and new burial customs spreading across Europe, and crucially, replacing the earlier Hunter-Gatherer societies. But exactly how this transformation happened was long debated. For decades, scholars argued between two main models. One said that farmers themselves migrated into Europe from Anatolia, bringing their technologies, weight of life and genetics with them. The other model said that Europe's Mesolithic Hunter-Gatherers learned farming techniques from their neighbours, adopting new tools and crops, while remaining the same people, genetically. And of course, there were people who said there was probably a bit of both. And by the 21st century, most scholars seemed to lean towards the idea that many Hunter-Gatherer groups adopted farming after encountering the migrating Neolithic farmers. And then, after about 2015, came the ancient DNA revolution. New techniques were developed to process whole genomes quickly and cheaply, meaning that hundreds of ancient individuals could be analysed and published in a series of groundbreaking papers. The data revealed that early European farmers were genetically distinct from the local Mesolithic Hunter-Gatherers. They were descendants of Anatolian farmers who multiplied and spread throughout Europe over generations. But in fact, it wasn't that simple. Anatolian farmers had mixed with the Mesolithic Hunter-Gatherers as they spread out from the Aegean across Europe. Now, one of the benefits of agriculture is that it allows a far greater population than is possible in most Hunter-Gatherer societies. And in Neolithic Europe, that meant the mixed groups were overwhelmingly Neolithic farmer with a small amount of Hunter-Gatherer admixture. For example, the Linear Pottery Culture, or LBK, after their German name Linear Band Keramik, of Central Europe, were almost all derived from Anatolian farmer ancestry but they also had an average of around 5% WHG ancestry. However, it looks like the development of the LBK as a distinct culture in around 5500 BC was stimulated by this interaction between the farmers and the local Hunter-Gatherers. Some of the earliest LBK individuals sampled had about 11% WHG ancestry, which is roughly the equivalent of having one WHG great-grandparent. But some early LBK people had 35% or even 50% WHG ancestry, which of course you would if you had one Hunter-Gatherer parent and one Neolithic farmer parent. And this mixing created a unique society still Neolithic farmers, but still distinct from the almost entirely Anatolian farmer Starcevo culture that came before it. Over time, the WHG ancestry, as I say, started to even out to around 5% in the general population of their LBK descendants. So really what I'm trying to say here is that even though the Hunter-Gatherer way of life in that region was extinguished by the arrival of the farmers, they also contributed culturally and genetically to the society that they were absorbed by. And I think this is important to remember that even long before the coming resurgence of WHG genes, the Neolithic farmers of Europe were also descendants of the Hunter-Gatherers who were here before them. And similar processes happened as Neolithic people expanded across the Mediterranean coasts, mixing to one degree or another with WHG populations and creating new cultures like the Cardial Wear culture. There were ongoing admixture events over the centuries, but these two different societies, the farmers and the Hunter-Gatherers, seemed to have had generally limited contact. Partly this was due to low population densities amongst the Hunter-Gatherers, but more so, perhaps, because they preferred different environments. The farmers sought river valleys with particular fertile soils, while the Hunter-Fisher foragers moved seasonally between resource-rich locations like coasts and wetlands, woodlands and uplands. Archaeology shows that there were often farmers and Hunter-Gatherers living relatively close to each other, but seemingly having little to no interaction in many cases. Maybe in some cases there was tension and conflict, and in others they just ignored each other. And this is the stuff that really fascinates me. You wonder about these thousands of interactions over the centuries. How these two different peoples met one another, what they thought of each other, and how they responded. Often there's only hints of these interactions. For example, domesticated pig bones might be found at a Mesolithic Hunter-Gatherer site. But whether they traded for those pigs as meat or living animals, or if they rustled them or stole them, or started to raise pigs themselves, is hard to say. It really fires your imagination. Genetic studies mainly show the occasional gene flow of Hunter-Gatherers into farming groups, but they can't reveal how this happened. We must assume there were some marriages and Hunters joining farming groups. No doubt some of this was voluntary, and other times was the result of intergroup violence. Like abductions following raids, sexual violence, forced assimilation, and maybe slavery, or some other kind of hierarchical inclusion and mixing. And we do see evidence for ongoing conflict in certain areas. At the limits of the LBK expansions in the North and West, some farmers built fortified enclosures. Judging by the wounds seen on skeletons, these were mainly to defend against raids by other farmers. But the density in the frontier zones suggests they were also built to protect from the Hunter-Gatherers beyond. The LBK territories in Central Europe did not extend all the way to the coasts of the North Sea and the Baltic. Along those coasts, the Hunter-Fisher foragers maintained higher population densities because of the rich maritime resources. And between the two societies was a kind of no-man's land where neither side lived. And this situation continued here for about a thousand years. After about 5000 BC, the LBK society collapsed for complex reasons and descended into an orgy of violence. There were massacres and a population decline. Settlement areas contracted and long-range trade and cultural contact was disrupted. Neolithic societies in the region became more differentiated and dissolved into distinct local cultures. And in Northwest Europe, the two strands of Neolithic farmers merged again as the cardio-ware groups of the Mediterranean pushed north through France and displaced or merged with post-LBK groups. And then around 4400 BC is when we start to see Hunter-Gatherer influence spread into some of these Neolithic cultures. And we see an increasing percentage of WHT ancestry and crucially, the replacement of many Neolithic farmer male lineages with Hunter-Gatherer ones. In Iberia, there were Mesolithic groups persisting especially in the Atlantic coastal region for 500 years after the Neolithic farmers arrived on the peninsula. And after about 4500 BC, there was an increase in Hunter-Gatherer ancestry in the Neolithic farmers. This is around the same time, 4400 BC as the proportion of WHT ancestry begins to rise again in Central Europe to between 15 to 40% and even over 50% in some individuals As well as overall ancestry, there are also increases in male and female WHT lineages. And this is perhaps the most significant point. The Y-Haplogroup evidence which traces male lineages, it's just a marker really passed on from father to son. This evidence shows the appearance of Haplogroups typical of Hunter-Gatherers in these farmer populations. The Y-Haplogroup does not in itself represent ethnicity. It's just a marker that shows lines of descent and can be useful in tracking migrations and population turnovers. Most of the Neolithic farmers before now had carried Y-Haplogroups like G2a from Anatolia. But these start to be replaced by WHG Haplogroups especially I2a. Perhaps 50% of the Anatolian male lineages in Europe are replaced by WHG ones. In some local cultures in North and West Europe, it was closer to 100%. While the Mitochondrial Haplogroups passed on by women to their children largely continue. But there are also some increases in WHG Mitochondrial Haplogroups. So this pattern shows the so-called resurgence, the increase in WHG genes in Neolithic farming societies was largely male biased. Now the Hunter-Gatherer resurgence didn't happen everywhere and long colonized places like Greece, the Balkans and Croatia likely didn't have many Hunter-Gatherers left around them. But with the increase in Hunter-Gatherer ancestry in Central Europe, new cultures emerged that began farming in new environments that had been formerly occupied by Hunter-Gatherers. These new cultures were a mixture of Central European Neolithic farmers who already had Ancient Hunter-Gatherer admixture and Mediterranean Neolithic farmers who also had Ancient Hunter-Gatherer admixture along with more recent admixture from local Hunter-Gatherers. These new societies were very successful and eventually expanded over generations all the way to the coasts of the Atlantic, the Baltic, the North Sea and the English Channel. After 4000 BC, some of these groups would also begin to colonize Britain and Ireland. These Neolithic farmers would once again out-compete and partially absorb the WHG who were living there. Significantly, every single Neolithic male who colonized Britain and Ireland apart from a couple of exceptions in Ireland carried the WHG I2AY haplogroup. But the real question is how did these late surviving Hunter-Gatherer groups manage to seemingly take over these Neolithic farming societies? On the face of it, this looks something like conquests of these farming groups carried out over a few generations where the WHG men slaughtered many of the Neolithic farmer men and took over their societies. Despite the genetic data, there is no clear evidence yet for the mass slaughter of Neolithic farmer men around this time. No massacre sites of men killed by WHG weaponry. And by the way, Neolithic men were no pushovers. They were well used to fighting each other with their maces, axes and adzes and bows and arrows. They had long built enclosures and fortified settlements to defend themselves from attack. So how could WHG conquer powerful settled societies like this? Well, perhaps these farmer groups had first suffered their own setbacks that weakened them. Perhaps issues like local overpopulation and resource exhaustion, disease epidemics and extreme competition for remaining resources leading to reciprocal genocidal attacks. A significant factor might be the climatic downturn seen from around 4500 BC which saw colder weather, soils becoming wetter and increased westerly winds and storminess around the North Atlantic. It could be population collapses weakened these societies and left them vulnerable to attacks from neighbouring hunter-gatherer groups. But it's interesting though that in this hypothetical scenario, the hunter-gatherer raiders didn't just abduct Neolithic farmer women and take them back to the Baltic coasts and wetlands and carried on being hunter-gatherers. Instead, they moved into these Neolithic communities and took them over, becoming farmers themselves. That's not at all an obvious decision especially if these were fragile societies. Genesisists point out that the replacement of many male lineages doesn't necessarily mean the men were all killed, only that they were out-competed in terms of reproduction by the incoming lineages. However, in practice, it's very hard to even imagine how that would work without violence and coercion. Nevertheless, perhaps this idea of violent conquests and takeovers isn't the whole story. For example, we could imagine a farming group weakened by internal conflicts and a neighbouring forager group weakened by disease who come to a mutual agreement to some kind of reciprocal support, maybe to defend each other from enemies, to share resources, to share territory and even exchange people in marriages. Perhaps in response to the deterioration of climatic conditions, the farmers suffered more than the hunter-gatherers did which gave the hunter-gatherers a competitive advantage. Maybe their skills and ancestral knowledge with resource gathering became extremely important when crops failed and herds perished. Another possibility, and all these hypotheses can be true at the same time, is that WHG groups thriving in woodlands, wetlands, uplands and coastal regions ultimately did adopt agriculture along with women from farming groups and adapted existing farming techniques in these new environments. The archaeological evidence shows in regions from Belgium to the Alps new kinds of landscapes previously inhabited by hunter-gatherers were being farmed for the first time after about 4400 BC. And in this era, new archaeological cultures emerge that genetic testing shows have this increased WHG ancestry and I2A YHAPA groups. Like the Mikkelsberg culture covering parts of modern Germany, France and Belgium. These people farmed new landscapes in new ways. Instead of living in large longhouses in long-term settlements like the LBK, manuring fields to maintain soil fertility, they seem to have taken up slash-and-burn agriculture across less fertile soils. People often lived in more ephemeral houses, clearing forests, farming crops for a few seasons until fertility declined and then moving on clearing more land. Cleared land and regrown secondary forest was then given over to the grazing of cattle changing the landscape from ancient oak and elm forests to forest pastures of hazel, alder, birch and ash. They and the other new cultures of Central Europe also built large fortified enclosures probably controlled by powerful elites along with huge burial monuments in stone and earth. The new cultures of Central Europe also expanded especially after the climate warmed again from 4000 BC. The Funnel Beaker culture descended from the Micklesburg culture expanded northwards into southern Scandinavia and the colonisation of Britain and Ireland began at the same time. They brought the increased WHG ancestry and I2A male lineages with them. Some have speculated that it was only this influx of hunter-gatherer culture that enabled the colonisation of these northern regions. The WHG people had thousands of years of experience navigating the waters of the Baltic and perhaps their boat building and maritime skills were the key to the Funnel Beaker expansions and the colonisation of Britain and Ireland. It's possible of course that these new societies were also speaking languages derived from the WHG people. It's not always the case historically when these mixing events happen but if the male lineages were WHG perhaps they also brought and passed on their hunter-gatherer languages too. Some people think the Neolithic megalithic culture the practice of building earth mounds standing stones and monumental megalithic tombs had its origins in Mesolithic societies. Perhaps it emerged from the mixing of southern European farmers central European farmers and local hunter-gatherers. Anyway, I think it's interesting to speculate that the megalith builders of sites like Stonehenge might have only been about 20% WHG by ancestry but they could have been speaking languages descended from those of their European Mesolithic ancestors. Starting after about 3000 BC these societies of Central and Western Europe would start to be changed and replaced by incoming cultures genes and lineages from Eastern Europe in the form of the Corded Ware culture bringing their Western step-herder ancestry with them. But it's worth pointing out that the Corded Ware also carried WHG admixture especially after mixing with Neolithic European societies like the Globular Amphora culture and there were later pulses of WHG ancestry and lineages into subsequent societies too like the migration from the Baltic that helped form the Unotecha culture in around 2200 BC. The classic WHG YHAPPA group I2A still continues especially concentrated in Central and Eastern Europe and people of European descent today still carry a proportion of Mesolithic hunter-gatherer DNA. I think it's worth reconsidering these old notions of a wave of Neolithic farmers starting in Anatolia and completely wiping out and replacing the Mesolithic inhabitants of Europe. Of course this was a devastating process for many individuals and groups who saw their lands taken by colonists probably in violence possibly after suffering from introduced infectious diseases and surely many of these early admixture events were not voluntary but likewise the Mesolithic people were not simply passive victims of this colonization and they were not wiped from existence. For generations they persisted in marginal lands and beyond the frontier maybe raiding, trading and learning from the farmers. Remember that these colonists already had partial hunter-gatherer ancestry and maybe that helped communication and understanding and then perhaps due to centuries of climatic changes they watched as the farmers declined and struggled and broke up into smaller groups and some hunter-gatherers maybe moved in and took advantage of their weakness taking over whole communities or maybe the two kinds of peoples joined together and came up with new approaches for how to live. Either way from the crisis powerful leaders emerged building new huge structures in the landscape to protect the living and to honor the worthy dead. These new societies then expanded and colonized lands that had never been farmed before and built tens of thousands of monuments many of which still exist today. So I don't see the story of Mesolithic Europeans as one of failure and extinction even if their way of life ultimately came to an end. Instead I think it's an incredible story of long resistance resilience adaptation and resurgence. Thanks again to MyHeritage for sponsoring this video. If you want to find out about your ancient ancestry and support my channel at the same time be sure to hit the link in the description or scan the QR code on screen and use my coupon code DAN25 to get your DNA kit for this limited time deal at this incredibly low price. Thank you very much indeed to my patrons and channel members for supporting my work. Now to find out more about the colonization of Neolithic Central Europe please watch this video on the LBK culture and their astonishing longhouses which were in 5000 BC the largest structures the world had ever seen. Thank you for watching.