Tag: Iron Age

Evolution of pigmentation in anatomically modern humans of Europe: a new paradigm?

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The colours of human skin, eyes and hair in living people across the world are determined by variants of genes (alleles) found at the same place on a chromosome. Since chromosomes are inherited from both mother and father, an individual may have the same two alleles (homozygous), or one of each (heterozygous). A dominant allele is always expressed, even if a single copy is present. A recessive allele is only expressed if the individual inherits two copies of it. Most characteristics of individuals result from the interaction of multiple genes, rather than a single gene. A commonly cited example is the coloration of eyes. If we had a single gene for eye colour – that of the iris – that had alleles just for blue (recessive or ‘b’) and one for brown (dominant or ‘B) pigmentation, brown-eyed individuals would have one or two ‘B’ alleles (bB or BB), whereas those with blue eyes would have to have two ‘blue’ alleles (bb). But inheritance is more complicated than that: there are people with green, hazel or grey eyes and even left- and right eyes of different colour. Such examples suggest that there are more than two genes affecting human eye colour, and each must have evolved as a result of mutations. Much the same goes for hair and skin coloration.

A group of scientists from the University of Ferrara in Italy have analysed highly detailed ancient DNA in anatomically modern human remains from Russia (Palaeolithic), Sweden (Mesolithic) and Croatia (Neolithic) to tease out the complexities of pigmentation inheritance. Then they applied a statistical approach learned from that study to predict the likely skin-, eye- and hair pigmentation in 348 less detailed genomes of ancient individuals whose remains date back to 45 Ma ( Silvia Perretti et al, 2025. Inference of human pigmentation from ancient DNA by genotype likelihood. Proceedings of the National Academy of Science, v. 122, article e2502158122; DOI: 10.1073/pnas.2502158122).

An artist’s impression of a Mesolithic woman from southern Denmark (credit: Tom Bjorklund)

All the hunter-gatherer Palaeolithic individuals (12 samples between 45 and 13 ka old) bar one, showed clear signs of dark pigmentation in skin, eyes and hair – the outlier from Russia was probably lighter. Those from the Mesolithic (14 to 4 ka) showed that 11 out of 35 had a light eye colour (Northern Europe, France, and Serbia), but most retained the dark skin and hair expected in descendants of migrants from Africa. Only one 12 ka hunter-gatherer from Sweden had inferred blue eyes, blonde hair, and light skin.  The retention of dark pigmentation by European hunter-gatherers who migrated there from Africa has been noted before, using DNA from Mesolithic human remains and in one case from birch resin chewed by a Mesolithic woman. This called into question the hypothesis that high levels of melatonin in skin, which protects indigenous people in Africa from cancers, would result in their producing insufficient vitamin D for good health. That notion supposed that out-of-Africa migrants would quickly evolve paler skin coloration at higher latitudes. It is now known that diets rich in meat, nuts and fungi – staple for hunter-gatherers – provide sufficient vitamin-D for health at high latitudes. A more recent hypothesis is that pale skins may have evolved only after the widespread Neolithic adoption of farming when people came to rely on a diet dominated by cereals that are a poor source of vitamin-D.

However, 132 Neolithic farmers (10 to 4 ka ago) individuals studied by Perretti et al. showed increased diversity in pigmentation, with more frequent light skin tones, yet dark individuals persisted, particularly in southern and eastern Europe. Hair and eye colour showed considerable variability, the earliest sign of red hair showing up in Turkey. Even Copper- and Bronze Age samples ( 113 from 7 to 3 ka) and those from Iron Age Europeans (25 from 3 to 1.7 ka ago) still indicate common retention of dark skin, eyes and hair, although the proportion of lighter pigmentation increased in some regions of Europe. Other analyses of ancient DNA have shown that the Palaeo- and Mesolithic populations of Europe were quickly outnumbered by influx of early farmers, probably from the Anatolian region of modern Turkey, during the Neolithic. The farming lifestyle seems likely to have allowed the numbers of those who practised it to rise beyond the natural environment’s ‘carrying capacity’ for hunter-gatherers. The former inhabitants of Europe may simply have been genetically absorbed within the growing population of farmers. Much the same absorption of earlier groups seems to have happened with the westward migration from the Ukrainian and Russia steppes of the Yamnaya people and culture, culminating in the start of the European Bronze Age that reached western Europe around 2.1 ka, The Yamnaya introduced metal culture, horse-drawn wheeled vehicles and possibly Indo-European language.

So the novel probabilistic approach to ancient DNA by Perretti et al. also casts doubt on the diet-based evolution of light pigmentation at high latitudes. Instead, pulses of large population movements and thus changes in European population genetics probably account for the persistence of abundant evidence for dark pigmentation throughout Europe until historic times. The ‘lightening’ of Europeans’ physiognomy seems to have been vastly more complex than previously believed. Early Europe seems to have been almost bewilderingly diverse, which make a complete mockery of modern chauvinism and racism. The present European genetic ‘melting pot’ is surprisingly similar to that of Europe’s ancient past.

Holocene migrations of people into Britain

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People assigned to a variety of human species: Homo sapiens – H. neanderthalensis (Swanscombe, 400 ka and several later times ) H heidelbergensis (Boxgrove, ca 500 ka, )H. antecessor (Happisburgh, ca 950 ka) – have left signs of their presence in Britain. Human occupancy has largely depended on climate. Around 9 times since the first known human presence here, much of Britain was repeatedly buried by glacial ice to become a frigid desert for tens of thousands of years. Between 180 and 60 ka only a couple of flint artefacts found in road excavations in Kent hint at Neanderthal visitors. For most of the Late Pleistocene the archipelago seems to have been devoid of humans. Arguably, Europe’s first known anatomically modern humans occupied several caves in Devon, Derbyshire and South Wales as early as around 43 ka, while climate was cooling, only to abandon Britain during the Last Glacial Maximum (24 to 18 ka ago). As climate warmed again thereafter, sporadic occupation by Late Palaeolithic hunter-gatherers occurred up to the sudden onset of the frigid Younger Dryas (12.9 ka). Once warming returned quickly 11,700 years ago, sea level was low enough for game and hunter gatherers to migrate to Britain; this time for permanent occupancy. Bones of the earliest known of these Mesolithic people have yielded DNA and a surprise: they were dark skinned and so far as we can tell remained so until the beginning of Neolithic farming in Britain around 6100 years ago. The DNA of most living Britons with pale skins retains up to 10% of inheritance from these original hunter gatherers.  Much the same is known from elsewhere in NW Europe. In the early Holocene it was possible to walk across what is now the southern North Sea thanks to Doggerland. Following a tsunami at around 8.2 ka this rich area of wetland vanished, so that all later migration demanded sea journeys.  

Mesolithic people remained in occupation of the British Isles for another two millennia. A wealth of evidence, summarised nicely in Ray, K. & Thomas, J. 2018, Neolithic Britain, Oxford University Press, suggests that there was a lengthy period of overlap between Mesolithic and Neolithic occupation around 4100 BCE. The main difference between the two groups was that Neolithic communities subsisted on domesticated grains and animals, while those of the Mesolithic consumed wild resources. Cultural clues in archaeological finds, however, suggest a lot in common, such as the erection of various kinds of monuments. Posts of tree trunks, sometimes arranged in lines, were raised in the Mesolithic and lines of probably ritual pits were dug. Both ‘traditions’ continued into the Neolithic and evolved to stone monuments, to which were added burials of different kinds. It is worth noting that Stonehenge was developed on a site that held much earlier, large totem-pole like posts, with a nearby spring that had hosted regular gatherings of Mesolithic people. Signs of Mesolithic occupation in Britain extend just as widely as do those of Neolithic practices. A study of DNA from 7 Mesolithic skeletons and 67 of early Neolithic age (Brace, S. and 20 others 2019. Population Replacement in Early Neolithic Britain. Nature Ecology & Evolution, v. 3, p. 765-771; DOI: 10.1038/s41559-019-0871-9) revealed that early Neolithic people did not wipe out the genetic make-up (either by complete displacement or annihilation) of their predecessors. About 20 to 30% of Neolithic DNA was inherited from them; as would be expected from assimilation of a probably much smaller number of hunter-gatherers into a larger population  of  immigrants who brought farming and herding from Asian Turkey (Anatolia). Such ‘hybrid’ genetics was widespread in Europe and they are referred to as the Early European Farmers (EEF). As Ray and Thomas suggest, aspects of Mesolithic culture may have been adopted by the newcomers across the British Isles from Orkney to Wiltshire.

Around 2400 BCE the earliest Neolithic ceremonial site at Brodgar on Orkney was destroyed to the accompaniment of an enormous feast that consumed several hundred cattle. At about the same time several men, whose tooth geochemistry indicated an origin in the European Alps, were buried on Salisbury Plain together with the earliest metal artefacts known from Britain (copper knives), the accoutrements of archery and distinctive, bell-shaped pottery beakers. Stonehenge was ‘remodelled’ shortly afterwards, with the addition of its giant trilithons, four of which were later adorned with carvings of metal axes and daggers. The Early Bronze (or Chalcolithic) Age had arrived! A 2018 study of ancient DNA from Bronze Age burials in Europe suggested a far more drastic swamping of Neolithic genetic heritage by the ‘Beaker people’ (Olalde, I. and a great many others 2018. The Beaker phenomenon and the genomic transformation of northwest Europe. Nature, v. 555, p. 190-196; DOI: 10.1038/nature25738). The skeletons from Britain analysed by Olalde et al. apparently suggested that, within a few hundred years, up to 90% of the Neolithic gene pool had been removed from the British population. Who were these people who used metals and the distinctive Bell Beakers, where did they come from and what did they do?

The closest match to the British and western European Bronze Age DNA was that associated with the Yamnaya people from the steppes of SE Ukraine and Southern Russia who had developed a culture centred on herding. They had also adopted the wheel from people of the Mesopotamian plains and had domesticated the horse for riding and pulling carts: ideal for their semi-nomadic lifestyle and for moving en masse. After 3000 BCE they spread into Europe, as widely recorded by their distinctive beakers and the presence of their DNA in the genomes of later Europeans. Their burials – in ‘kurgans’ – resembled the round barrows that appeared on Salisbury Plain and elsewhere during the Bronze Age. The DNA replacement data from 2018 were limited and held few clues to how it happened. One possibility for such a dramatic change could be a violent takeover that drove down the population of British Neolithic people. To address the broader influence of migration in more detail and over a loner time span, a team led by the Universities of York and Vienna, and Harvard Medical School (Patterson, N. and a great many others 2021. Large-scale migration into Britain during the Middle to Late Bronze Age. Nature, early online release; DOI: 10.1038/s41586-021-04287-4) used ancient DNA from 793 individuals excavated in Britain (416 individuals) and continental Europe (377) from Bronze- to Iron Age sites (2300 to ~100 BCE).

The proportion of Early European Farmers DNA in British individuals from the Bronze Age (2400 BCE) to the Iron Age (750 BCE to 43 CE). Note the ‘fuzzy’ nature of the data, and that the decline in EEF in British individuals was not as great as earlier analyses had shown. Remarkably, the ‘Amesbury Archer’, who brought the first metals to Britain, had a higher proportion of EEF ancestry than the Early Bronze-Age average. (Credit: Patterson et al. Fig. 3)

The new data from Britain suggest that the migrants, who crossed the Channel later in the Bronze Age, were of mixed ethnicity, but most carried EEF genes. The influence of earlier migrants from the Yamnaya heartlands is present, but so too are relics of Mesolithic ancestry. Interestingly, the British data show a much larger increase in the genes associated with lactase persistence, which marks the ability of adults to digest milk, than was apparent in the wider European population (50% compared with about 7% in Eastern Europeans of the time). Whatever the impact of the first influx of metal-using people – it may have been culturally decisive in Britain – by the end of the Bronze Age the EEF ‘signature’ had increased in peoples’ genomes. Rather than some kind of invasion, the influx was more likely to have been a sustained movement of people to Britain over several hundred years By the Iron Age, almost half the ancestry of Britain, particularly in England and Wales, was once again predominantly of EEF origin (around 40% of the mixture), but culture had become completely different. There are even suggestions that the influx brought with it the beginnings of Celtic languages. Yet the data leave a great deal of further analysis to be undertaken.

See also: Drury, S.A. 2019. Genetics and the peopling of Britain: We are all hybrids, People and Nature; Ancient DNA Analysis Reveals Large Scale Migrations Into Bronze Age Britain, SciTechDaily, 28 December 2021.