Throughout the history and prehistory of Europe, the average height of its inhabitants has gone up and down many times. The reasons for this are many. Some of the more obvious examples being diet, climate, population movements (and thus the changing of the gene pool) as well as selection.

Height is a polygenic trait. That means it’s controlled not just by one gene, but by multiple different genes. Height has a high heritability and genetic height is fairly easy to measure.There may or may not be more undiscovered genes other than the ones we have thus far identified that are associated with increased stature. So, while the exact accuracy of height prediction may not be spot-on, we can still get a decent grasp of what kind of genetic height potential different ancient populations had. Especially when supplemented with archaeology, which allows us to take a look at the actual skeletons of ancient humans to gain a better understanding of their physicality. The current scientific consensus is that around 80% of final height is determined by genetics, while the remaining 20% is due to environmental factors.

In present-day Europe, we can observe a significant variation in national average height. Some of it is related to living standards and dietary practices, but the maximum genetic height potential of a population is mainly determined by DNA. It can be said that there is a North-South gradient of height variation in Europe, although there are some exceptions. So what is it that makes the Portugese average height differ so from the Dutch average height? The answer may lie in prehistory.

Abbreviations

UP = Upper Paleolithic

LGM = Late Glacial Maximum

WHG = Western Hunter-Gatherer

EHG = Eastern Hunter-Gatherer

WSH = Western Steppe-Herder

AHG = Anatolian Hunter-Gatherer

ANF= Anatolian Neolithic Farmer

Height of Paleolithic, Mesolithic, Neolithic and Bronze Age Europeans

Many populations have come and gone in Europe. Groups that physically differed drastically from each other. Broadly speaking, it can be said that Upper Paleolithic Europeans were very tall. Mesolithic Europeans were significantly shorter, but still had a high in-population variation with some individuals being very short and others being quite tall. Neolithic European farmers, especially in Southern Europe, had a very low average height. In the Bronze Age we can observe a dramatic rise in height again, but it wasn’t until the early 2000s that some European populations returned to a similar average height as the Upper Paleolithic European hunter-gatherers. Let’s take a more detailed look at the height of some ancient European groups.

Upper Paleolithic European hunter-gatherers (Aurignacians, Gravettians)

Upper Paleolithic Europeans were hunter-gatherers living in an extreme and brutal climate. Aurignacians are believed to have been some of the first groups (or even the first) of anatomically modern humans to enter Europe. Physically, the Cro-Magnon of this culture were extremely robust and also tall for their time, with most male skeletons being around the 179 cm mark, or just below 5´11. The Aurignacian culture was later replaced by the Gravettian, and the skeletons from this shift are different. This (along with a clear change in material culture) led archaeologists to believe that there was a population replacement, or at least some change, and in modern times this seems to have been vindicated through DNA research. Y-DNA samples from the Aurignacians are sparse, but they seem to have been carriers of haplogroup C-V20. Recent results from Hajdinjak et al. (2021) revealed one sample of F-M89 and two additional samples carrying C1. The main haplogroup of the Gravettians seems to have been I-M170 (I*) and genetically, the expansion of the Gravettian archaeological industry is associated with the spread of hunter-gatherers carrying mainly haplogroup I-M170 and the Vestonice-cluster autosomal DNA profile. The Gravettians were also fairly robust, although less strongly built than the Aurignacians. In terms of stature, they were one of the tallest ancient populations to ever have existed. The average Gravettian male stood around 182 cm, or just under 6´.

There are many theories about what made the Gravettians so tall. For starters, it’s safe to assume that one factor that likely contributed to their impressive stature was that they hunted megafauna. This would have allowed for a massive intake of daily calories and a diet very rich in protein and fat. The latter would have been especially vital for survival in the European climate at the time. Mammoths, reindeer and large wolves provided the Gravettians with energy-rich nutrition. In addition to large game hunting, their advanced tools and hunting tactics allowed them to supplement their diet with smaller animals that they caught using nets. The Gravettians on the coastlines of Europe were known to hunt sea animals as well. This well-rounded, nutritious and high caloric diet likely allowed for a near ideal environment for physical growth. Some research has suggested that the hunter-gatherers of the Gravettian selected for stride length (long legs, shorter upper body) due to this trait benefitting their highly mobile lifestyle.

Western hunter-gatherers (WHG)

Mesolithic Europeans had a mean height of 166 cm for males. This is a rather dramatic height difference when compared with Upper Paleolithic Gravettians, or even the Aurignacians. With the extinction of Mammoths in their part of Europe, the WHGs did not have access to megafauna. The WHGs mainly hunted deer and boar. On the coasts there were thriving communities of hunter-fishers. It’s possible that in this post-LGM environment that offered less diverse options for game, the hunter-gatherers were unable to acquire such a high amount of daily calories as the Gravettians were. Another factor to take into account is selection. WHGs were less mobile than Gravettians, and generally had more geographically limited hunting territory. They certainly moved around, but they didn’t trek across different parts of Europe in the same way that the Gravettians often did. That may have contributed to less selection for the long-legged builds that were so common among Gravettians.

Of course, it’s also possible that the differences in height can mainly be attributed to an admixture event with another incoming population to Europe, for example from Anatolia. It was suggested in Feldman et al. 2019 that a hunter-gatherer population living in the Balkans during the Mesolithic (Iron Gates HG) may have been something intermediate between WHG, EHG and a third Anatolian/Near Eastern hunter-gatherer component. This indicates that WHGs had contacts with AHG and AHG-like populations quite a long time before Anatolian Neolithic Farmers migrated to Europe. Keep in mind, WHG are in no way a direct continuation of Gravettians. Gravettian ancestry in WHGs amounted to about 37%.

These things considered, WHGs still had a fairly high polygenic height score. This indicates that they had the genetic potential to become tall, althouh perhaps not quite as tall as their (paternal) Gravettian ancestors. It’s possible that for some reason related to environmental stress, the WHGs were unable to reach their genetic height potential. Straus, 1995 showed that climate degradation during the LGM resulted in an increase in population density in Europe, which in turn led to an overexploitation of resources. This caused the hunter-gatherers of Europe to widen the breadth of their diet. Protein sources that may not even have been considered options by the Gravettians may have been consumed by WHGs who were unlikely to have considered them low-rank or low-quality food, simply due to their lack of options. Small aquatic protein sources is one such example. As a whole, it is definitely possible that selection for reduced body size as a means to reduce nutritional demands started among the Solutreans (who suceeded the Gravettians) and was completed among Magdalenians (who suceeded Solutreans and preceded WHGs) and among WHGs. It’s worth mentioning that these hardy Mesolithic hunter-gatherers of Europe were still significantly taller than the Early European Farmers, which was likely a result of diet, lifestyle and genetics.

Yamnaya steppe pastoralists

DNA samples from the Yamnaya culture show high polygenic height scores and display signals of selection for elevated height. One thing that pastoralist populations (both prehistoric, historical and modern) have in common is that they tend to be taller than their more sedentary neighbours. Being a mix of 2 populations with high predicted genetic height (EHG, CHG) as well as having a diet and lifestyle that was near-ideal for high stature, it’s no wonder that these people became so tall.

The average male height of skeletons associated with the Yamnaya cultural horizon is 174 cm. It is fully possible that the rather impressive height of the Yamnaya herders and autosomally related steppe cultures was one of many contributing factors to their success, and the subsequent spread of haplogroups R1a and R1b across Europe. It’s worth mentioning that the CHG admixture in Yamnaya may be very old and not from a more recent admixture event in the Neolithic. It’s possible that a CHG-like population had been living on the steppe since the Mesolithic.

A study by Marnetto et al. (2022) examined ancestral genomic contributions to complex traits in the modern Estonian population. What they found was that Western Steppe Herder (or Yamnaya-related) ancestry is strongly associated with increased stature. It is worth keeping in mind that their results may not translate over to every other European population, however, as selection and various local ancestry components also play a role in regards to height.

In any case, Yamnaya-related admixture correlates quite well with increased average height in present-day Europe. However, there are certainly exceptions to this, which we will get to later in this article.

Scandinavian hunter-gatherers (SHG)

Another Mesolithic population with a very high polygenic height score. It doesn’t come as a shock, given that the SHGs were in a similar position as Yamnaya/WSH in that they were a mix of 2 hunter-gatherer populations (WHG + EHG in the case of SHGs) with high polygenic height scores. Additionally, SHG skeletons generally show signs of excellent health. This was a thriving group of people, and it’s blatantly reflected in their skeletal remains and genomes.

Cardium Pottery/Cardial Ware culture farmers

EEF as a whole display lower polygenic height scores than hunter-gatherer populations.

Southern European EEF groups seem to differ from Central/Northern European EEF groups when it comes to genetic height. This can likely be explained by the significantly lower WHG input in Southern EEF along with different dietary practices. EEF groups in Central and Northern Europe would have had a diet higher in protein than the more southern groups. This was because they supplemented their agricultural lifestyle with hunter-gathering to a greater extent. Adding to that, there is evidence suggesting that the EEF groups in Central and Northern Europe may have consumed dairy products. This would have made their diets more rich in protein and calories than the more southern farmers.

Selection comes in, as always. In Neolithic and Chalcolithic Iberia, we can actually observe a significant selection for reduced height. At least if we’re to believe the results of Mathieson 2015. The study found that Iberian Neolithic and Chalcolithic farmers had a lower polygenic height score than both early Anatolian farmers and Middle Neolithic farmers in Germany, Poland and Scandinavia.What exactly it was that caused this is hard to say, but it’s possible that it had to do with famines. Studies like Berbesque et al, 2014 have shown that early farmers in Europe starved much more often than hunter-gatherers did. It’s possible that these continuous famines simply culled tall individuals from the gene pool and served as a selection against tall stature. It would have been unadvantageous to be tall when calories were sparse since tall individuals need more daily calories to sustain themselves, and a tall stature is not necessarily helpful in acquiring nutrition as a farmer. As a hunter-gatherer, however, tall stature is often an advantageous trait. What we see in the case of EEF groups in Southern Europe, is that there was a population with a low polygenic height score and thus a low genetic height potential that also seems to have gone through selection for reduced height. This resulted in a population with a low average height.

Inconsistencies and contradictions

A study by Berg et al. (2019) could not find evidence of selection for reduced height in Neolithic and Chalcolithic Iberia. As such, their results seem to contradict those of Mathieson and colleagues (2015). Their results did however suggest that height variation in Europe depends on variation in hunter-gatherer ancestries. High amounts of WHG, EHG and CHG admixture can be traced to an increased polygenic height score and thus, an increased genetic “height potential” of a population, according to them. That seems to add up with the results of other studies, but the relation between HG admixture and increased height doesn’t seem as clear-cut as they want to make it seem.

For example, Grasgruber et al, 2019, proposes that the modern Albanian population is significantly shorter (8.9cm shorter) than Montenegrins independent of nutrition due to the fact that the Albanian population has much lower degrees of WHG and Yamnaya-related admixture than the Montenegrins. What they fail to mention is that Greeks have slightly lower Yamnaya-related and WHG admixture than Albanians, but are still much taller on average, and the Greek male average height of 177 cm is on par with many populations in Western Europe. Another example of where the prediction that European height should follow Yamnaya-related admixture does not seem to apply is if you compare, let’s say Norwegians, and Montenegrins. Norwegians have a very high percentage of Yamnaya-related admixture, and quite a lot of WHG admixture too. Montenegrins on the other hand, don’t have particularly high steppe nor WHG admixture, but still have a significantly greater average height than Norwegians. This clearly shows that it might not always be as simple as Yamnaya + WHG = taller. Rather, it seems that more recent selection likely also plays an important role in determining the height of a modern population, not just admixture from prehistoric groups.

This isn’t the only example of recent studies being somewhat inconsistent when discussing the topic of genetic height. Another study by Grasgruber from 2014 focused on examining the role of nutrition and genetics on average heights of European populations. The study argues that since populations with unusually tall males align remarkably well with the distribution of haplogroups I1, R1b-U106 and I2a1b-M423 that could be explained by the fact that the genetic predisposition for unusually tall stature was already present among Upper Paleolithic hunter-gatherers (Gravettians).

What the study fails to address is that just because a population happens to have high frequencies of Gravettian Y-DNA, that doesn’t mean that they have much total DNA from Gravettians. In fact, actual percentual genetic ancestry from Gravettians among modern Europeans is very low. That should come as no surprise, given how many other groups have come and gone since the time of the Gravettians. The study makes a few other errors that are more easy to look past given that the study is now 7 years old, such as suggesting that the reason for why the Iberians and the Irish have lower average heights is because many males of those populations carry R1b-S116, which the study associates with the Magdalenians and declares native to Western Europe. Of course, nowadays we know that this is not the case, and Magdalenians did not carry R1b-S116. What Grasgruber et al. 2014 should have addressed is that the Y-chromosome makes up only a miniscule percentage of the full genome of a person, and does not have any direct effect on height or appearance in general. The results of the study could better be explained by the fact that present-day carriers of I1 and R1b-U106 tend to have very high levels of Yamnaya-related ancestry on the autosomes, as well as high levels of lactose tolerance and a recent history of high living standards, all 3 factors likely contributing significantly to the high average heights of these populations. In the case of I2a1b-M423 which is common in the Balkans, a reasonable explanation for why some South Slavic populations are so tall might be due to a case of strong, recent selection for increased height.

In May 2022, Grasgruber published another study that mainly touches on what his previous studies on the topic of height and Y-DNA haplogroups discussed. The authors of Grasgruber et al. (2022) conclude that the high average height of men from the Adriatic coast and the Dinaric Alps is due to having a high frequency of haplogroup I-M170. While it is an interesting hypothesis, Grasgruber and colleagues once again miss out on the autosomal aspect of things, as well as on the possibility of recent selection.

As such, Grasgruber’s data is valuable in itself but both the older and the new studies lack the kind of nuance we would require to really reach the same conclusions as he and his team did. As explained earlier, the differences in height between European populations can be far more accurately explained by different gradients of autosomal components, living standards and selection.

Conclusions

Genetic height is complicated. While there undoubtedly exists a North-South gradient of height variation in present-day Europe, this variation can not be explained simply through varying percentual degrees of admixture from prehistoric groups. However, current research indicates that admixture from ancient hunter-gatherer populations of Europe and the Caucasus may be a decent predictor of a high polygenic height score. Ancient genomes suggest that hunter-gatherer populations were not only genetically predisposed to being tall, but also often had ideal living conditions for physical development. There is also substantial evidence of some European populations having gone through strong selection pressure for increased stature in relatively recent times.

References:

Grasgruber, Pavel (2019): The coast of giants: an anthropometric survey of high schoolers on the Adriatic coast of Croatia https://www.ncbi.nlm.nih.gov/pmc/articles/PMC6475134/

Mathieson, Ian (2015): Genome-wide patterns of selection in 230 ancient Eurasians https://www.ncbi.nlm.nih.gov/pmc/articles/PMC4918750/

Berg, Jeremy et al (2019): Polygenic Adaptation has Impacted Multiple Anthropometric Traits https://www.biorxiv.org/content/10.1101/167551v3.full.pdf

Rosenstock, Eva (2019): Human stature in the Near East and Europe ca. 10,000–1000 BC: its spatiotemporal development in a Bayesian errors-in-variables model https://link.springer.com/article/10.1007/s12520-019-00850-3

Mathieson, Ian (2015): Eight thousand years of natural selection in Europe https://www.biorxiv.org/content/10.1101/016477v2

Holt (2008): Hunters of the Ice Age: The Biology of Upper Paleolithic People https://onlinelibrary.wiley.com/doi/pdf/10.1002/ajpa.20950

Berbesque (2014):Hunter–gatherers have less famine than agriculturalists https://www.ncbi.nlm.nih.gov/pmc/articles/PMC3917328/

Cox, Samantha (2019): Genetic contributions to variation in human stature in prehistoric Europe https://www.pnas.org/content/116/43/21484

Feldman (2018): Late Pleistocene human genome suggests a local origin for the first farmers of central Anatolia https://www.biorxiv.org/content/10.1101/422295v1

Grasgruber, Pavel (2014): The role of nutrition and genetics as key determinants of the positive height trend https://www.sciencedirect.com/science/article/pii/S1570677X14000665

Tornberg, Anna (2015): A tale of the tall : A short report on stature in Late Neolithic–Early Bronze Age southern Scandinavia https://portal.research.lu.se/ws/files/3913084/7853683.pdf

Hajdinjak, Mateja (2021): Initial Upper Palaeolithic humans in Europe had recent Neanderthal ancestry https://www.nature.com/articles/s41586-021-03335-3

Marnetto, Davide (2022): Ancestral genomic contributions to complex traits in contemporary Europeans https://www.cell.com/current-biology/fulltext/S0960-9822(22)00108-7

Pavel, Grasgruber (2022): Mapping the Mountains of Giants: Anthropometric Data from the Western Balkans Reveal a Nucleus of Extraordinary Physical Stature in Europe https://www.mdpi.com/2079-7737/11/5/786/htm

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