Genetic history of the British Isles
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Population research using DNA is initiating research into the genetic history of the British Isles. Genetically, the population native to the British Isles is closely associated with the larger region of Western Europe, and in particular with the European "Atlantic fringe" characterized for example by Y chromosome (patrilineal) haplogroup R1b1b2 (R-M269). This haplogroup originated less than 18,000 years ago (possibly much less) in Western Asia, and may have entered Europe in the Neolithic.
Immigration to Great Britain during the British Empire, including immigration in recent decades has further changed the picture, with the "White British" population accounting for 87 percent in the United Kingdom Census 2001.
The study of geographical patterns in genetic types became a mainstream and frequently cited science in the late 1970s. It gave a method of trying to understand past human migrations. Prior to this various other research methods were used for similar effect, such as examining the distributions of blood type. Later published studies used mitochondrial DNA to study the female line of descent, and then it became possible to use Y chromosome DNA to study male descent lines. More recently, use has been made of technologies which can test hundreds of thousands of possible mutation points (SNPs) in the rest of the human genome (the autosomal DNA).
Y and mtDNA haplogroups
Bryan Sykes published in 2001 a study of European mitochondrial DNA (mtDNA) in which he identified the seven principal haplogroups in Europe.. He subsequently set up the Oxford Genetic Atlas Project, which has collected more than 50,000 DNA samples, 10,000 of which were obtained in Britain and Ireland. He produced an analysis of 6000 of these and has published it in his book Blood of the Isles using twelve mitochondrial haplogroups and five Y-DNA haplogroups for various regions of the Isles. He has given his conclusions separately for Ireland, Scotland, Wales and England. Sykes gives useful paternal and maternal clan distribution maps, (website is no longer available) as well as maintaining a web site where the detailed data is given. His principal conclusion is that the paternal and maternal settlement patterns are different for the British Isles.
Stephen Oppenheimer used the Weale and Cappelli study data, with that of Rosser et al. for Europe, in new analyses of the data published in The Origins of the British. He uses phylogeographic mapping to produce many distribution diagrams, covering Europe as well as Britain and Ireland. His principal conclusion is that there are both east and west coast entry routes to Britain and Ireland. Walter Bodner is currently undertaking a further survey of the DNA of the Isles but only a preliminary TV programme has so far been produced. In Ireland population genetic studies have been undertaken by a team under Dan Bradley, including surname studies. Databases on Britain and Ireland, as well as on various surnames, are also being built up from personal DNA tests, for example at FTDNA.
Sykes and Oppenheimer have each given nicknames to various haplogroups to allow easier recognition, including the principal ones in the Isles. The following names are those of Sykes unless otherwise stated:
- H Helena (Sykes), Helina (Oppenheimer)
- I Isha
- J Jasmine
- T Tara
- U Europa (Oppenheimer)
- U2 Uta
- U3 Uma
- U4 Ulrika
- U5 Ursula
- V Velda (Sykes), Vera (Oppenheimer)
- W Wanda
- X Xenia
- R1b Oisin (Sykes), Ruisko (Oppenheimer)
- I Wodan (Sykes), Ivan (Oppenheimer)
- R1a Sigurd (Sykes), Rostov (Oppenheimer)
- E3b Eshu
- J Re
Oppenheimer has broken down R1b into sixteen founding clusters in the Isles of which the more important ones are: -8 Rob (Frisian Modal Haplotype), -9 Rox (Basque), -10 Ruy (Atlantic Modal Haplotype, called Atlantis by Sykes), -14 Rory (Gaelic Modal Haplotype). He has also broken down haplogroup I into three, of which the most important ones for the Isles are I1a (Ian) and I1c (Ingert).
Paleolithic Europeans seem to have been a homogenous population, possibly due to a population bottleneck (or near-extinction event) on the Iberian peninsula, where a small human population is thought to have survived the glaciation, and then expanded into Europe during the Mesolithic period. The assumed genetic imprint of Neolithic incomers is seen as a cline, with stronger Neolithic representation in the southeast of Europe and stronger Paleolithic representation in the west of Europe. There are thought to have been three separated pockets of human habitation in Europe during the last major glaciation (the end of the Paleolithic and the Pleistocene), on the Iberian peninsula, in the Balkans and north of the Black Sea in what is now the Ukraine. The Y chromosome haplogroups from these populations are thought to correspond to R1b (Iberian), I (Balkans) and R1a (Ukraine); these three haplogroups occur all over Europe, but their frequencies are not spread uniformly.
These early genetic investigations have shown that the biological influence of post-mesolithic immigration on Britain may have been smaller than previously thought, marked more by continuity than change. The Oxford archaeologist David Miles, in The Tribes of Britain, states that 80% of the genetic makeup of native Britons probably comes from "just a few thousand" nomadic tribesmen who arrived 12,000 years ago, at the end of the Ice Age. This suggests that later waves of immigration may have been too small to have significantly affected the genetic homogeneity of the existing population. However, Miles himself acknowledged that the techniques used to explore genetic ancestry are still in their infancy and that many more samples are needed to fully understand the origins of the British people.
Stephen Oppenheimer has recently argued that neither Anglo-Saxons nor Celts may have had much impact on the genetics of the inhabitants of the British Isles, and that British ancestry can mostly be traced back to ancient peoples similar to the modern-day Basques instead. Current estimates of the genetic contribution of Anglo-Saxon migrants range from less than 10,000 to as many as 200,000. A recent study by a team from the Department of Biology at UCL based on computer simulations indicate that an apartheid-like social structure in early Anglo-Saxon England provides a plausible explanation for a high-degree of continental male-line ancestry in England.
The above article, based on analysis of the Y chromosomes of men currently living in the United Kingdom, the Western Isles, Orkney, Shetland, Friesland, Denmark, North Germany, the Republic of Ireland, Norway and the Basque Country, is "consistent with the presence of some indigenous component in all British regions". For the sake of this study samples from the Basque Country were considered indigenous (a putative paleolithic Y chromosome). These studies cannot distinguish between Danish, Frisian and German (Schleswig-Holstein) Y chromosomes. Within England areas with the highest concentration of Germanic (Danish-Viking/Anglo-Saxon) Y chromosomes occurred in areas associated with the Danelaw and Danish-Viking settlement, especially York and Norfolk. In these areas, about 60% of Y chromosomes are of Germanic origin. It should be noted that this indicates an exclusively male component. The extent of Danish/Anglo-Saxon contribution to the entire gene pool of these areas is also dependent on the migration of women. For example, if it is assumed that few or no Germanic women settled in these areas, then the Germanic contribution to the gene pool is halved to 30%, and in turn if greater numbers of women did settle, the contribution could be even higher than 60%.
In two recently published books (Blood of the Isles by Bryan Sykes and The Origins of the British by Stephen Oppenheimer) both authors state that according to genetic studies (mainly Haplogroup R1b as well as E3b, I, J (Y-DNA), and mtDNA), most Britons descend mainly from ancient populations of the Iberian Peninsula, as a result of different migrations that took place during the Mesolithic and the Neolithic, which laid the foundations for the present-day populations in the British Isles, indicating an ancient relationship among the populations of Atlantic Europe. Both authors claim that there is evidence for this on both the Y-chromosomes and maternal DNA. Stephen Oppenheimer asserts that he is even able to date when migrations to the British Isles took place and where exactly within the Iberian peninsula these different migrations originated. Stephen Oppenheimer also claims that there were neolithic invasions to Scotland from Norway prior to Norwegian Viking invasions and that the Vikings would take the same invasion routes as the previous Neolithic invaders.
In Origins of the British (2006), Stephen Oppenheimer states (pages 375 and 378):
- By far the majority of male gene types in the British Isles derive from Iberia (Spain and Portugal), ranging from a low of 59% in Fakenham, Norfolk to highs of 96% in Llangefni, north Wales and 93% Castlerea, Ireland. On average only 30% of gene types in England derive from north-west Europe. Even without dating the earlier waves of north-west European immigration, this invalidates the Anglo-Saxon wipeout theory …
- … 75-95% of British Isles (genetic) matches derive from Iberia … Ireland, coastal Wales, and central and west-coast Scotland are almost entirely made up from Iberian founders, while the rest of the non-English parts of the British Isles have similarly high rates. England has rather lower rates of Iberian types with marked heterogeneity, but no English sample has less than 58% of Iberian samples …
In page 367 he also states in relation to Rossers's pan-European genetic distance map:
- In Rosser's work, the closest population to the Basques is in Cornwall, followed closely by Ireland, Wales, Western England, Scotland, Spain, Belgium, Portugal, East Anglia and then northern France"
Bryan Sykes, in his book Blood of the Isles (2006), states (p. 280):
- … the presence of large numbers of Jasmines’s Oceanic clan, says to me that there was a very large-scale movement along the Atlantic sea board north from Iberia, beginning as far back as the early Neolithic and perhaps even before that. The number of exact and close matches between the maternal clans of western and northern Iberia and the western half of the Isles is very impressive, much more so than the much poorer matches with continental Europe.
- The genetic evidence shows that a large proportion of Irish Celts, on both the male and female side, did arrive from Iberia at or the same time as farming reached the Isles.
- The connection to Spain is also there in the myth of Brutus … This too may be the faint echo of the same foundation myth as the Milesian Irish and the connection to Iberia is almost as strong in the British regions as it is in Ireland.
- Here again, the strongest signal is a Celtic one, in the form of the clan of Oisin, which dominates the scene all over the Isles. The predominance in every part of the Isles of the Atlantic chromosome (the most frequent in the Oisin clan), with its strong affinities to Iberia, along with other matches and the evidence from the maternal side convinces me that it is from this direction that we must look for the origin of Oisin and the great majority of our Y-chromosomes. The sea routes of the atlantic fringe conveyed both men and women to the Isles.
Complicating matters in analysing the data has been the fact that Danish Viking and Anglo-Saxon are at present genetically indistinguishable, making it uncertain which group has had a larger impact on the current population.
On the other hand, a team of geneticists led by Santos Alonso does not observe any particular link between Basques and Celtic populations beyond that provided by the Paleolithic ancestry common to European populations.
Cro-Magnons are known to have had a presence in Great Britain by 27,000 years ago, due to the discovery of the skeletal remains of the "Red Lady of Paviland". During the Last Glacial Maximum (LGM)) around 27,000 years ago Northern Europe may have been completely depopulated. Humans probably returned to the region of the British and Irish peninsula about 14,700 years ago, as glaciers started to retreat, after an absence of about 10,000 years. Around 9500 BC rising sea levels due to ice melting caused Ireland to become separated from Britain, while around 6500 BC Britain became separated from continental Europe.
About 4000 BC the "Neolithic Revolution" reached Britain and Ireland, with domestication of animals, arable farming and pottery. Again a new invasion was postulated. A population "wave of advance" was proposed  but this now seems to have had only a minor affect on the Isles. Francis Pryor estimates that by 4000 BC the population of Great Britain was around 100,000 while that of Ireland was some 40,000. For 2000 BC his estimates are 250,000 and 50,000.
Celtic and Anglo-Saxon invasions
The Celtic and Anglo-Saxon invasions of the British Isles are difficult to document genetically. Sykes' and Oppenheimer's books, based upon recent genetic studies, particularly those of Weale etc. and Capelli etc., make a case for the majority of Britons having ancestors from the Iberian Peninsula, as a result of a series of migrations that took place during the Mesolithic and, to a lesser extent, the Neolithic eras. Oppenheimer and Sykes believe that during these periods there was also a smaller immigration across the North Sea. They also believe that in the total immigration in the historical period may only be 5%.
Sykes sees little genetic evidence relating to people from the heartland of the Hallstatt and La Tène cultures. On the paternal side he finds that the "Oisin" (R1b) clan, which is in the majority, has strong affinities to Iberia, with no evidence of a large scale arrival from Central Europe. He considers that the genetic structure of Britain and Ireland is "Celtic":
|“||if by this we mean descent from people who were here before the Romans and who spoke a Celtic language.||”|
Oppenheimer's theory is that the modern day people of Wales, Ireland and Cornwall are mainly descended from Iberians with lesser input from Germany and Scandinavia. Furthermore, Oppenheimer claims that Celtic split from Indo-European earlier than previously suspected, some 6000 years ago, while English split from Germanic before the Roman period. Oppenheimer believes that a Germanic language that become English was spoken by the tribes of what is now England long before the arrival of the Anglo-Saxon.
In Origins of the British (2006), Stephen Oppenheimer states (pages 375 and 378):
|“||By far the majority of male gene types in Britain and Ireland derive from Iberia (modern Spain and Portugal), ranging from a low of 59% in Fakenham, Norfolk to highs of 96% in Llangefni, north Wales and 93% Castlerea, Ireland. On average 30% of gene types in England derive from north-west Europe. Even without dating the earlier waves of north-west European immigration, this invalidates the Anglo-Saxon wipeout theory …|
… 75-95% of Britain and Ireland (genetic) matches derive from Iberia … Ireland, coastal Wales, and central and west-coast Scotland are almost entirely made up from Iberian founders, while the rest of the non-English parts of Britain and Ireland have similarly high rates. England does have lower rates of Iberian types with marked heterogeneity, but no English sample has less than 58% of Iberian samples …
A study was made of 100 English people by Piercy et al. and published in 1993, the data from which has been incorporated into later studies. In 2001 a team led by Jim Wilson published a paper on the differing male and female roles during cultural transmission in the British Isles. A paper published in 2002 by Michael Weale of University College London focussed on the Anglo-Saxon invasion. They sampled seven small towns across central England and North Wales in comparison with samples from Friesland and Norway, to look for evidence of immigration from the continent. They examined three population processes: simple splitting with subsequent divergence, single mass migration, and continuous background migration. They produced a genetic distance map that showed significant differences between the English and Welsh samples, with the Friesland sample clustering with the former. They concluded that mass Anglo-Saxon migration was the most likely event, by default. However, they did not consider that Friesland and England may have had similar immigration histories prior to this, while there were special circumstances relating to one of their sites. A later paper by Weale and others considered how this situation could have arisen by a process of apartheid. A later larger study of 25 small towns in England was made by a team led by Cristian Capelli and the results published in 2003. A genetic distance map was included in their paper, which contained data from Wales, Scotland and Ireland as well as Norway and North Germany/Denmark. In contrast to the Weale study, this paper showed a gradual change between western and eastern England. They concluded that there had not been complete population replacement anywhere in the British Isles. They said that the most surprising conclusion was the limited continental input in southern England.
The percentage of the R1b haplotype on the Y chromosomes of English males, at about 64%, indicates that they may be descended primarily from the earliest Paleolithic peoples thought to have recolonised western Europe from a western Ice Age refuge after the end of the last major glaciation some 25-15 thousand years ago.
Y chromosome analysis of men from Britain, Denmark, Ireland, Germany, Norway, Friesland and the Basque Country of Northern Spain and South Western France has revealed that the Germanic (Danish/North German/Frisian) component in the male line of descent is higher in some areas of England than others. It is highest in York and Norfolk, where the Germanic Y chromosome occurs in about 60% of men, while indigenous Y chromosomes comprise about 40%, and lowest in the west of the country. The research cannot distinguish between Danish (the presumed source of Danish-Viking settlers to East and Northern England), North German (Schleswig-Holstein, modern era) and Frisian (Anglo-Saxon) Y chromosomes. It concludes "these data are consistent with the presence of some indigenous component in all British regions". Also, this research cannot make reference to the extent of settlement by Anglo-Saxon/Danish-Viking women. Therefore even in places like York, the total genetic contribution of these peoples would be less than 60% if fewer women than men migrated, and conversely it would be greater if more women than men settled.
Bryan Sykes found that the maternal clan (haplogroup) pattern was similar throughout England, but with a definite trend from east and north to the south and west. There were slight variations in the Helena clan with more striking differences in the Jasmine clan. He says that there were two branches of the latter (the Ocean and Land ones) that came via the Atlantic coast and Baltic respectively. The minor clans are mainly found in the east of England.
Computer simulations have shown that it is theoretically possible for a small Anglo-Saxon population that was politically and economically dominant to support larger families, which in turn could have resulted in a faster population growth for the dominant class. This model has been likened to apartheid in South Africa. These data assume that there is a 50% Anglo-Saxon Y chromosome occurrence through England, though this assumption has been contested by more recent genetic surveys. In some areas, notably Cornwall (and to a lesser extent Cumbria and Lancashire), some people claim a stronger ethnic connection to the ancient Britons (see Wales below), consequently some nationalists claim that Cornish people are distinct from English people.
Genetic research has demonstrated that, in the Scots populations sampled, the R1b marker is common. R1b averages 75% in Scotland, although east Scotland has lower levels of R1b. This marker is also common in most populations of Western Europe reaching highest frequencies in the west of Ireland and in the Basque Country of northern Spain and southern France.
In two recently published books (Blood of the Isles by Bryan Sykes and The Origins of the British by Stephen Oppenheimer) both authors state that according to genetic studies (mainly Haplogroup R1b as well as E3b, I, J (Y-DNA) and mtDNA), most Scots, along with the rest of the population of the British Isles descend mainly from ancient populations of the Iberian Peninsula, as a result of different migrations that took place during the Mesolithic and the Neolithic, which laid the foundations for the present-day populations in the British Isles, indicating an ancient relationship among the populations of Atlantic Europe. Both Authors claim that there is evidence for this on both the Y chromosomal and maternal DNA. Stephen Oppenheimer is even able to date when migrations to the British Isles took place and where exactly within the Iberian peninsula these different migrations originated. Stephen Oppenheimer also claims that there were neolithic invasions to Scotland from Norway prior to Norwegian Viking invasions and that the Vikings would take the same invasion routes as their previous neolithic invaders.
In the Northern Isles, Sykes estimated that there were 42% of men with Viking ancestry and 37% in Shetland. He found remarkable similarities between the mtDNA of the Northern Isles and that of Norway, suggesting that the Vikings brought their women with them and that their settlement had been peaceful. In the "Pictish" areas of Grampian and Tayside the mtDNA was little different from the rest of Britain. On the west coast of Scotland it was found that 15% of the DNA was Norse in origin for both types of DNA. However, there were 22% of males but only 11% of females with Norse origin in the Hebrides, decreasing to 7% and 2% repectively in Argyll. In Argyll it appears that there was a 30-40% replacement of Pictish by Gaelic Y chromosomes. Sykes found in the Hebrides evidence of clans that are spread all along the Atlantic fringe, in the form of Mediterranean Jasmines and the younger Taras.
The consensus in Wales today is that they regard themselves as Celtic, claiming a heritage back to the Iron Age tribes, which themselves, based on modern genetic analysis, would appear to have had a predominantly Paleolithic and Neolithic indigenous ancestry. When the Roman legions departed Britain around 400, a Romano-British culture remained in the areas the Romans had settled, and the pre-Roman cultures in others.
Genetic evidence suggests that most Welsh people descend from the population of an ancient Paleolithic western Ice Age refuge, which was situated in northern Spain and southern France, roughly surrounding the Pyrenees. As the result of different migrations that took place during the Mesolithic and Neolithic, these laid the early foundations for the present-day populations in the British Isles and may indicate an ancient relationship between the populations of Atlantic Europe. According to Stephen Oppenheimer 96% of lineages in Llangefni in north Wales derive from Iberia. Genetic research on the Y-chromosome has shown that Welsh men, like Irish men share many lineages with men from the Basque Country of northern Spain and south-western France, Welsh men may have more descent from Neolithic people than either Irish or Basque men.  Haplogroup R1b averages from 83-89% amongst Welsh men.
The pattern of maternal clans was found by Sykes to be very similar to that of Ireland, with that of Helena predominating. A few differences were found between north, mid and south Wales. There was a closer link between north and mid Wales that either had with the south. In the Y-DNA there was practically no sign of Viking settlement but there was some Wodan that appeared to have been there for a long time. The Oisin clan is very common, reaching 86% in mid Wales, and the Atlantis type is a greater proportion than in Ireland. The diversity of the Oisin clan was found to be low, which usually suggests a recent arrival date. However, other indicators suggest that mid Wales has been among the most stable and longest settled of any region in the Isles.
Current genetic research supports the idea that the Y chromosomes and mtDNA of people living in Ireland are mainly descended from the indigenous European Paleolithic (Old Stone Age hunter gatherers) population, with a smaller Neolithic (New Stone Age farmers) input particularly with the Y chromosomes and mtDNA from people of the Celtiberians or Galicians of Galicia (Spain). Y chromosome analysis also seems to indicate that the Vikings that settled in Dublin came from Norway rather than Denmark. The frequencies of Y chromosome haplotypes in the Irish population are similar to that of most other populations of Atlantic Europe, especially the Basques of northern Spain and southern France.Mitochondrial DNA, which is inherited through the female line, shows part of the maternal ancestors of the Irish to be of broad north European origin, while confirming the Atlantic link.
There is genetic evidence that a majority of Irish maternal and paternal lineages descend from Paleolithic and Mesolithic west Europeans who arrived after the end of the last Ice Age, noting the high presence of the genetic marker known as Atlantic Modal Haplotype (AMH) among the Irish. This is associated with haplogroup R1b and is the most common R1b haplotype in western Europe. R1b averages 90% amongst Irish Y chromosomes, and 90% amongst Y chromosomes of Basque people. The AMH+1 haplotype has a frequency of 53% in central Ireland (Castlerea) and 60% in the Basque country, it also achieves a greater than 50% frequency in other regions of the British Isles, namely Pitlochry, Oban, Morpeth, Penrith, Isle of Man, Llangefni, Haverfordwest, Midhurst and Cornwall. Curiously, the Irish samples of Castlerea and Rush (Dublin) have some of the highest frequencies of haplogroups R1 and R1a1 (37% and 45% respectively), with the Scottish samples of Durness and Stonehaven being the only others that surpass 30%.
Bryan Sykes found that 10% of the Irish population were direct descendants of Ursula but most had modified mtDNA. He calculated a date of 7300 BC for their entry into Ireland. Similar dates were found for the other clans, long before the Iron Age Celts were supposed to have arrived. Little difference was found between the maternal clans in the four provinces. However, he did find differences in their Y-DNA clans, with Oisin going from 73% in Leinster to 98% in Connacht. It was found that in the Oisin clan there was a greater proportion of men with Gaelic names than Anglo-Norman ones.
Geneticists have found that seven men with a rare Yorkshire surname carry a genetic signature previously found only in people of African origin. All the men tested positive for haplogroup A1, a Y chromosome genetic marker which is highly west African specific. Haplogroup A1 is extremely rare and has only ever been found 25 times, again only in people of African origin. Haplogroup A1 is a subclade of Haplogroup A which geneticists believe originated in Eastern or Southern Africa.
The individuals had no knowledge of any African heritage in their family. The researchers hypothesized that the presence of this haplogroup in Yorkshire could stem from the recruitment of Africans for the construction of Hadrian's Wall by the Romans or result from intermarriage with an African slave, some of whom rose quite high in society.
Geneticists have shown that former American president Thomas Jefferson, who was of Welsh descent, along with two other British men out of 85 British men with the surname Jefferson carry the rare Y chromosome marker K2 which is typically found in East Africa and the Middle East. Haplogroup K2 is extremely rare in Europe but phylogenetic network analysis of its Y-STR (short tandem repeat) haplotype shows that it is most closely related to an Egyptian K2 haplotype. The presence of scattered and diverse European haplotypes within the network is nonetheless consistent with Jefferson's patrilineage belonging to an ancient and rare indigenous European type.
- ^ Balaresque et al. (2010), "A Predominantly Neolithic Origin for European Paternal Lineages", PLoS Biol. 8 (1), doi:10.1371/journal.pbio.1000285, PMID PMC2799514, http://www.ncbi.nlm.nih.gov/pmc/articles/PMC2799514
- ^ B. Arredi, E. S. Poloni and C. Tyler-Smith (2007). "The peopling of Europe". in Crawford, Michael H.. Anthropological genetics: theory, methods and applications. Cambridge, UK: Cambridge University Press. p. 394. ISBN 0-521-54697-4.
- ^ Karafet, TM; Mendez, FL; Meilerman, MB; Underhill, PA; Zegura, SL; Hammer, MF (2008). [Expression error: Missing operand for > "New binary polymorphisms reshape and increase resolution of the human Y chromosomal haplogroup tree."]. Genome research 18 (5): 830–8. doi:10.1101/gr.7172008. PMID 18385274.
- ^ ISOGG 2009
- ^ Cavalli-Sforza, L.L. (2001) Genes, People and Languages. London: Penguin.
- ^ Sykes, B. (2006). Saxons, Vikings, and Celts: The Genetic Roots of Britain and Ireland. London UK: Transworld Publishers. ISBN ISBN 978-0393330755 ISBN 0393330753.
- ^ Sykes, Sykes, The Seven Daughters of Eve, 2001,
- ^ Sykes has since found that an eighth is also common.
- ^ Gibbons (2000)
- ^ Review of The Tribes of Britain James Owen, National Geographic 19 July 2005.
- ^ Stephen Oppenheimer, "Myths of British ancestry", Prospect, October 2006, accessed 21 September 2006.
- ^ PDF (206 KiB)
- ^ a b "A Y Chromosome Census of the British Isles"; Cristian Capelli, Nicola Redhead, Julia K. Abernethy, Fiona Gratrix, James F. Wilson, Torolf Moen, Tor Hervig, Martin Richards, Michael P. H. Stumpf, Peter A. Underhill, Paul Bradshaw, Alom Shaha, Mark G. Thomas, Neal Bradman, and David B. Goldstein Current Biology, Volume 13, Issue 11, Pages 979-984 (2003). Retrieved 6 December 2005.
- ^ Special report: 'Myths of British ancestry' by Stephen Oppenheimer | Prospect Magazine October 2006 issue 127
- ^ a b Celts descended from Spanish fishermen, study finds - This Britain, UK - Independent.co.uk
- ^ From the Cover: Genetic evidence for different male and female roles during cultural transitions in the British Isles
- ^ 030705U491
- ^ European Journal of Human Genetics - Abstract of article: The place of the Basques in the European Y-chromosome diversity landscape
- ^ Stringer (2006), p. 182
- ^ Mithen (2003), p.120
- ^ see Renfrew, 1987
- ^ How did pygmy shrews colonize Ireland? Clues from a phylogenetic analysis of mitochondrial cytochrome b sequences
- ^ http://www.omniglot.com/blog/?p=516
- ^ Oppenheimer, S. (2006). The Origins of the British: A Genetic Detective Story: Constable and Robinson, London. ISBN 978-1-84529-158-7.
- ^ Sykes (2006), p.290.
- ^ The western refuge encompassed southern and eastern France, the Basque country and the north coast of Spain. (Oppenheimer, p. 103). In the Basque country R1b represents 90% of the male population. It is the only R haplogroup to have emerged from the western refuge after the last Ice Age. (Oppenheimer, p. 116).
- ^ a b c d e f Capelli et al. (2003)
- ^ Thomas et al. (2006).
- ^ Weatherhill.
- ^ Atlas of the Human Journey - The Genographic Project
- ^ Estimating the Impact of Prehistoric Admixture on the Genome of Europeans - Dupanloup et al. 21 (7): 1361 - Molecular Biology and Evolution
- ^ Estimating the Impact of Prehistoric Admixture on the Genome of Europeans - Dupanloup et al. 21 (7): 1361 Figure 04 - Molecular Biology and Evolution
- ^ Special report: 'Myths of British ancestry' by Stephen Oppenheimer, Prospect Magazine October 2006 issue 127
- ^ From the Cover: Genetic evidence for different male and female roles during cultural transitions in the British Isles, Proc Nat Ac Sci 2001, Apr 4. 98(9) 5078-5083
- ^ What happened after the fall of the Roman Empire?: BBC Wales-History. Retrieved 03 October 2006.
- ^ McEvoy et al. (2004)
- ^ a b Hill et al. (2000)
- ^ Wilson et al. (2000).
- ^ a b Yorkshire family linked to Africa
- ^ National Geographic Haplotype Map
- ^ Tracing Past Human Male Movements in Northern/Eastern Africa and Western Eurasia: New Clues from Y-Chromosomal Haplogroups E-M78 and J-M12
- Gibbons, Anne. (2000) Evolutionary Genetics: Europeans Trace Ancestry to Paleolithic People Abstract, Science: Vol. 290. no. 5494, pp. 1080–1081. Retrieved 15 February 2007.
- Hill, C. Origins of the English
- Hill, Emeline, W., Mark A. Jobling, Daniel G. Bradley. (2000) Y-chromosome variation and Irish origins PDF. Nature, Vol 404,. Retrieved 30 December 2005.
- Miles, David. The Tribes of Britain,
- Mithen, Steven 2003. After the Ice: A Global Human History 20,000-5000 BC. Phoenix (Orion Books Ltd.), London. ISBN 978-0-7538-1392-8
- McEvoy, Brian., Martin Richards, Peter Forster, and Daniel G. Bradley (2004) The Longue Durée of Genetic Ancestry: Multiple Genetic Marker Systems and Celtic Origins on the Atlantic Facade of Europe Am. J. Hum. Genet., 75:693-702, 2004. Retrieved 15 February 2007.
- Oppenheimer, Stephen 2006. The Origins of the British: A Genetic Detective Story. Constable and Robinson. ISBN 1845291581.
- Rosser, Zoe et al.. 2000. Y-chromosomal diversity in Europe is clinal and influenced primarily by geography. Americal Journal of Human Genetics 67, 1526-43.
- Stringer, Chris. 2006. Homo Britanicus. Penguin Books Ltd., London. ISBN 978-0-713-99795-8.
- Sykes, Bryan. 2001. The Seven Daughters of Eve
- Sykes, Bryan. 2006. The Blood of the Isles. Bantam Press. ISBN 0593056523
- Thomas, Mark G., Michael P. H. Stumpf and Heinrich Härke (2006) PDF (206 KiB) Proceedings of the Royal Society Published online. doi:10.1098/rspb.2006.3627. Retrieved 15 February 2007.* Wilson, James F., Deborah A. Weiss, Martin Richards, Mark G. Thomas, Neil Bradman, and David B. Goldstein (2001) PDF Proceedings of the National Academy of Science. 98: 5078–5083 . Retrieved 15 February 2007.
- The study headed by Dr Bradley was published in the American Journal of Human Genetics. Geneticists find Celtic links to Spain and Portugal