The Origin of Icelanders

England was invaded by Vikings from both Denmark and Norway. The influence of the Danish Vikings can be seen most strongly in Y chromosomes from York and Norfolk in the eastern regions that bore the brunt of the Danish invasions. The Norwegian Vikings operated to the north of the Danes. In the ninth century AD they captured the Orkney Islands to the northeast of Scotland and made them a base of operations. Norn, a form of old Norse, was spoken on the islands until the eighteenth century. Norwegian Vikings have left a strong genetic signature among Orcadians, as Orkney Islanders are known, but their traces can also be seen farther afield, particularly in Iceland.

From their base in Orkney the Norwegian Vikings sailed around the northern coast of Scotland and down the waterway between Britain and Ireland, making settlements on both the British and Irish sides. In AD 870, the Vikings discovered Iceland, several days' sail to the northwest of Scotland. Apart from some Irish hermits, who quickly left, the island was uninhabited. News of this virgin territory, with no hostile natives, soon got around, and for 60 years there was a steady stream of settlers. Immigration ceased in AD 930, perhaps because many of the trees had been chopped down, prompting an ecological crisis, and there was no more unclaimed farmland left. The island was then essentially closed to new immigration until modern times.

Iceland's genetic history has received much attention, both for its intrinsic interest as an isolated human population and because its population has become a leading source for discovering the genetic roots of common diseases ranging from cancer and heart disease to asthma and schizophrenia. These diseases are thought to result from several errant genes acting in combination. The errant genes are very hard to detect because each makes only a small contribution to the overall disease. For various reasons, including an excellent system of medical records, Iceland offers many advantages in searching for such genes. In 1996 Kari Stefansson, an American-trained Icelandic neurologist, put together a high powered genetic analysis company, DeCode Genetics, which has enjoyed considerable success in identifying disease genes in Icelanders and other populations. The company and its large pharmaceutical partners hope to develop diagnostic tests and drugs on the basis of the Icelandic findings. It is therefore of considerable interest to know if Icelanders are genetically similar enough to other populations, particularly those of the United States and Europe, for discoveries about their patterns of genetic disease to be relevant elsewhere in the world.

Icelandic records from the twelfth and thirteenth centuries, notably documents known as the Book of Settlements and the Book of Icelanders, indicate that although Norse Vikings directed the immigration to Iceland, the inflow included people from the Norse settlements in the Orkneys and the coastal regions of Scotland, northern England and Ireland. Most of these Norse invaders, after the initial conquest, had intermarried with the local population. Assuming these Vikings brought their families, many if not most of the women in the founding Icelandic population would have been British or Irish, and in either case of Celtic origin. The Book of Settlements mentions only a small proportion of the founding settlers by name but of those whose ancestry is recorded, only 5% of the men came from the British Isles but 17% of the women. In addition, the Vikings captured slaves in raids in both countries, many of whom were probably women.

Icelandic historians have developed the case that their country was probably founded by men who were mostly Norse and women who were mostly from the

British Isles, especially Ireland. This claim of descent from two important peoples, the Vikings and the Celts, helped to differentiate the Icelanders of the nineteenth and twentieth centuries from their much-resented rulers, the Danes. "The result of this conflation is the dominant modern concept of Icelandic origins, one that fuses the nobility and heroism of the Norse with the literary and other cultural traditions of the Irish and other peoples of the 'Celtic fringe,'" write a group of Icelandic and other experts.304

Given the power of modern genetics to deconstruct complex populations like that of England, it should be a simple problem to analyze the genetics of Iceland and check the validity of the historians' position. But it's not so easy. Comparison of Icelandic Y chromosomes with those in Scandinavia and the British Isles confirms that most of the male founders were indeed Norse, though not overwhelmingly so: some 20 to 25% of Iceland's founding fathers appear to have had Gaelic, meaning Celtic, ancestry, with the rest being of Norse 305

origin.

The founding mothers are much harder to trace. The patterns of mitochondrial DNA found among Icelanders today look generically European but without greatly resembling those of any particular country.306 They look, well, Icelandic. The reason is probably genetic drift, the random gain or loss of genetic signatures between generations, accentuated by the violent fluctuations in Iceland's population since the settlement. The Black Death killed 45% of the population in 1402-1404. A smallpox epidemic in 1708 reduced the population by 35%. Famine, the aftermath of a volcanic eruption, caused a 20% decline in 1784-1785. After each of these population declines and expansions, the characteristic mix of mitochondrial DNA signatures would have changed, pushing the population farther down a separate path from that of its source populations.

With the genetic answers being so Delphic, a team of researchers has resorted to the old technique of craniometry. They measured Icelandic skulls from the settlement period and compared them with medieval skull collections from Ireland and Norway. Unfortunately the Icelandic skulls were not in good enough condition to tell their sex. Overall, they seemed very similar to the Norse skulls and less like those of Ireland. The researchers say that "although our results do not preclude a significant Irish or other contingent among the settlers of Iceland, we conclude that the founding population was of predominantly (60-90%) 307

Norwegian origin."_

To help in its quest for disease genes, DeCode Genetics has assembled a genealogical database of the Icelandic population that extends back 1,100 years into the past. It is based on calfskin documents that hold the first 300 years of records, on church archives, and on the data from three complete censuses that were held starting in 1703. DeCode's genealogist, Thordur Kristjansson, reckons the database includes the names of about half the Icelanders who have ever lived, including 85% of those born in the nineteenth century and almost everyone who lived in the twentieth century. The database has enabled DeCode researchers to explore the historical dynamics of a human population in fascinating detail.

One finding is that generation times are shorter in mother-to-daughter lines of descent than in father-to-son lineages. The average interval between generations was 29 years in female lineages going back to 1698, 32 years for male lineages. The difference presumably reflects the simple fact that women tend to be younger than their husbands.

A greater surprise is how many people in one generation leave few or no descendants in the next. DeCode has traced the ancestry of all 131,060 Icelanders born from 1972 to 2002 back to two cohorts of ancestors. Of all contemporary Icelandic women born since 1972, 92% are descended from only 22% of the women born in the 1848-1892 cohort, and 86% of contemporary men are the progeny of just 26% of this group.

The progeny pyramid narrowed even more steeply going back to an earlier generation of ancestors, those born between 1698 and 1742. Because of the incompleteness of the genealogy for earlier centuries, the pedigree of many contemporary Icelanders could not be traced that far back. Nevertheless, DeCode researchers found that just 7% of the women born in the early eighteenth century period are the ancestresses of 62% of contemporary women, while 10% of the men of this period fathered 71% of contemporary males.309 Most people, in other words, have lines of descent that eventually go extinct, at least in a population the size of Iceland's, while just a few ancestors give rise to the majority of subsequent population cohorts.

This difference in reproductive success seems to be due largely to genetic drift, the force of which depends on the size of the population. Iceland's population fell to a post-settlement low of 33,000 after the 1708 smallpox epidemic but steadily increased from the beginning of the nineteenth century to its present level of 290,000. Even during this expansion, the influence of genetic drift was still at work. Before the end of the Pleistocene, there may have been many human populations no bigger than this, offering much grist for drift to work on.

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