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The Ancestors of the English

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Postby yialousa1971 » Sat Mar 26, 2011 9:15 pm

Excavating Past Population Structures by Surname-Based Sampling: The Genetic Legacy of the Vikings in Northwest England

The genetic structures of past human populations are obscured by recent migrations and expansions and have been observed only indirectly by inference from modern samples. However, the unique link between a heritable cultural marker, the patrilineal surname, and a genetic marker, the Y chromosome, provides a means to target sets of modern individuals that might resemble populations at the time of surname establishment. As a test case, we studied samples from the Wirral Peninsula and West Lancashire, in northwest England. Place-names and archaeology show clear evidence of a past Viking presence, but heavy immigration and population growth since the industrial revolution are likely to have weakened the genetic signal of a 1,000-year-old Scandinavian contribution. Samples ascertained on the basis of 2 generations of residence were compared with independent samples based on known ancestry in the region plus the possession of a surname known from historical records to have been present there in medieval times. The Y-chromosomal haplotypes of these 2 sets of samples are significantly different, and in admixture analyses, the surname-ascertained samples show markedly greater Scandinavian ancestry proportions, supporting the idea that northwest England was once heavily populated by Scandinavian settlers. The method of historical surname-based ascertainment promises to allow investigation of the influence of migration and drift over the last few centuries in changing the population structure of Britain and will have general utility in other regions where surnames are patrilineal and suitable historical records survive.

A useful test case to evaluate the potential of surname-based ascertainment is provided by the 1,000-year-old Viking settlement of the British Isles. The Viking age of raiding, exploration, trading, and colonization began in the late eighth century, with a series of attacks on the coasts of Britain, Ireland, and France. In England, a shift from raiding to permanent settlement began after AD 865. The modern consensus has moved from the earlier view of a mass migration of Viking settlers to a more geographically variable and gradual process, with much assimilation of local culture, under the administration of a Scandinavian elite (Richards 2004). Even though Viking rule in England came to an end nearly 1,000 years ago and the settlers were soon integrated linguistically and culturally, abundant evidence of Scandinavian influence remains today. Important archaeological findings exist, but the most striking evidence is linguistic and onomastic—many words in Standard English and in local dialects, and many place-names, are of Scandinavian origin. In some areas of English counties that formed part of the Danelaw (fig. 1), the region under the administrative control of the Vikings from the late ninth century, up to 70% of major place-names are in this category, with endings such as “-by” and “-thorp(e).”

In one version of events, Vikings of Norwegian origin, under their leader Ingimund, arrived in the region in AD 902 after having been expelled from Dublin. Æthelflæd, Lady of the Mercians, granted them land in north Wirral, where they settled (Wainwright 1942, 1948, 1975). More complex models have also been proposed, based on place-names analysis, including elements of migration from the Isle of Man as well as Dublin (Fellows-Jensen 1992). Specific historical evidence for the migration of Vikings into West Lancashire is lacking, although place-names and archaeological evidence clearly demonstrate that this occurred.

Widespread migration and population expansion in recent centuries present a particular problem in genetic analyses of mainland regions of Britain. The Wirral experienced a major influx of people since the medieval period, its population growing over 70-fold between 1545 and 1921 (Roberts 2002)—almost 10 times the national average increase. In conventional sampling strategies, where 2 generations of residence in an area are sufficient to qualify a DNA donor for participation in a study, the signal of Viking influence is likely to be weakened by the noise of more recent population movement

Where comparisons can be made, the proportions of Scandinavian admixture that we estimate differ somewhat from those seen in a previous study (Goodacre et al. 2005): for Shetland, we observe 41% Scandinavian ancestry compared with the previously published figure of 44% and the corresponding figures for Orkney are 50% compared with 31%. Such differences may reflect sampling variance or differences in the compositions of the parental and hybrid samples and in the marker resolution—we used greater numbers of both binary markers and microsatellites. Note that direct comparisons with the study from which we draw our comparative data (Capelli et al. 2003) are problematic because it used a different admixture method and considered admixture between a Norwegian sample and a sample chosen to represent Anglo-Saxons. The lowest proportions of Scandinavian admixture among our samples are seen in Llangefni and Mid-Cheshire, at 10% and 21%, respectively. In agreement with previous results (Capelli et al. 2003), a higher proportion (39%) is seen in the Isle of Man, where there is a known history of Viking presence, and Penrith (37%), which shows Scandinavian dialect influence (Reaney 1927). The modern samples from Wirral and West Lancashire both show 38% Scandinavian admixture, markedly higher than the nearby sample from Mid-Cheshire (21%). This is consistent with the historical and place-name evidence for greater Viking presence in Wirral and West Lancashire than in Mid-Cheshire.

The medieval sample from West Lancashire shows an increased proportion (51 ± 4%) of Scandinavian ancestry compared with its modern counterpart (38 ± 4%); the equivalent values for Wirral are 47 ± 5% and 38 ± 3%. These differences, revealed by our different sampling strategies, are likely to reflect a change in haplotype frequencies due to postmedieval immigration and are supported by genetic distances (FST) between the Norwegian sample and the Wirral and West Lancashire samples. FST between Norwegians and the West Lancashire modern sample is 0.130, whereas the value for the medieval sample is only 0.069; corresponding values for the Wirral samples are 0.162 and 0.096.

The surnames that we used to ascertain the medieval samples belong to a wide range of frequencies—from “Otty,” with only 146 bearers in 1998 (www.spatial-literacy.org/UCLnames/) to “Brown,” with 242,765 (supplementary table, Supplementary Material online). The more frequent names are likely to have had multiple founders and are relatively widespread in Britain and so may provide less reliable links to medieval presence in the specific regions under study. To address this, we subsampled from the 2 medieval samples by removing surnames with frequencies of greater than 20,000 (supplementary table, Supplementary Material online), resulting in reduced sample sizes of 26 and 30 for Wirral and West Lancashire, respectively. We then repeated the admixture analysis. In both cases, the proportions of Scandinavian admixture in the subsampled medieval populations increased further compared with the original medieval samples: admixture proportions in the modern and medieval subsampled Wirral populations are now 38 ± 3% compared with 51 ± 6% and the corresponding values for West Lancashire are 38 ± 4% compared with 53 ± 5%.


There you go dog..................


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Postby yialousa1971 » Sat Mar 26, 2011 9:17 pm

Summary:

Percentage of Scandinavian admixture in modern populations (average with other studies cited) : Anglesey (10%), Western Scotland (15%), Mid-Cheshire (21%), Western Isles and Skye (22.5%), Cumbria (37%), Wirral and West Lancashire (38%), Isle of Man (39%), Orkey (40%), Shetlands (42.5%).

Comparison with medieval samples : Wirral (47%), West Lancashire (51%)
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Postby yialousa1971 » Sat Mar 26, 2011 9:32 pm

Y Chromosome Evidence for Anglo-Saxon Mass Migration

Michael E. Weale,*1 Deborah A. Weiss,†1 Rolf F. Jager,*‡ Neil Bradman,* and
Mark G. Thomas*
*The Centre for Genetic Anthropology, Departments of Biology and Anthropology, University College London,
University of London; †Department of Anthropology, University of California, Davis; and ‡Faculteit Biologie,
Vrije Universiteit, Amsterdam, The Netherlands
British history contains several periods of major cultural change. It remains controversial as to how much these
periods coincided with substantial immigration from continental Europe, even for those that occurred most recently.
In this study, we examine genetic data for evidence of male immigration at particular times into Central England
and North Wales
. To do this, we used 12 biallelic polymorphisms and six microsatellite markers to define highresolution
Y chromosome haplotypes in a sample of 313 males from seven towns located along an east-west transect
from East Anglia to North Wales. The Central English towns were genetically very similar, whereas the two North
Welsh towns differed significantly both from each other and from the Central English towns. When we compared
our data with an additional 177 samples collected in Friesland and Norway, we found that the Central English and
Frisian samples were statistically indistinguishable
. Using novel population genetic models that incorporate both
mass migration and continuous gene flow, we conclude that these striking patterns are best explained by a substantial
migration of Anglo-Saxon Y chromosomes into Central England (contributing 50%–100% to the gene pool at that
time
) but not into North Wales.

Introduction
Following depopulation during the last glacial maximum
and subsequent resettlement by hunter-gatherers
ca. 7000 B.C.
, the history of Britain has been marked by
a series of cultural transitions. These include the appearance
of sedentary agricultural communities (the
Neolithic transition) (ca. 4000 B.C.), the arrival and
spread of Late Bronze-Iron Age and Celtic material culture
(ca. 1000–100 B.C.), Roman occupation and influence
(A.D. 43–410), the rise of Anglo-Saxon language
and culture (ca. A.D. 400–800), Viking invasions and
influence (ca. . 800–1000), and the Norman Conquest

(A.D. 1066) (Kearney 1989; Hunter and Ralson 1998;
Davies 1999).


Several patterns emerge from analyses of genetic
distance and population differentiation
(table 4 and fig.
3). Firstly, little genetic differentiation exists among the
Central English towns.
The only significant difference
in haplotype frequencies occurs between the neighboring
towns of Fakenham and North Walsham (P 5 0.032
for individual pairwise comparison or P 5 0.035 for
combined test of all English towns). Although more hg1
and fewer hg2 chromosomes were observed in the Midlands
than in East Anglia (table 2), Mantel tests on the
genetic versus geographic distance correlation and linear
regression on hg1 and hg2 frequencies reveal no significant
within-England clinal patterns.
Secondly, in contrast to the Central English towns,
the two North Welsh towns show highly significant differences,
both from each other and from the five Central
English towns
. Llangefni has a very high frequency of
hg1, tightly clustered around the modal haplotype (haplotype
#1 in table 3; haplotype 1.15 in Wilson et al.
2001) that has been found at high frequency in other
Atlantic populations, including Ireland (Hill, Jobling,
and Bradley 2000) and the Orkneys (Wilson et al. 2001).
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Postby yialousa1971 » Sat Mar 26, 2011 9:45 pm

Both Llangefni and Abergele have very low frequencies
of hg2 and dispersed haplotypes within this haplogroup,
a pattern which is consistent with these haplotypes entering
the North Welsh populations through separate, infrequent
admixture events.
Thirdly, no significant differences in haplotype frequencies
exist between Friesland and any of the Central
English towns. Comparisons between Norway and the
Central English towns, on the other hand, are all significant,
apart from Bourne (P 5 0.237), which may be
explained by the small number of samples collected
from this town (n 5 12). Furthermore, bootstrap tests
on RST values revealed that the Central English (all five
towns combined) are significantly more closely related
to the Frisians than they are to the North Welsh (Llangefni:

P , 0.001; Abergele: P 5 0.046) or to the Norwegians
(P 5 0.005). Both Friesland and Norway are
significantly different from the North Welsh towns. Similar
results were obtained using FST values based on haplogroup
frequencies, but tests on FST values based on
haplotype frequencies were not significant because of
the large number of singletons at this level. Taken together,
these results suggest considerable male-line commonality
between Central England and Friesland.

Wilson et al. (2001) identified two haplotypes—(1)
2.47 (haplotype #60 in our table 3), and (2) 3.65 (haplotype
#107 in our table 3)—that proved useful in inferring
a Viking contribution to the Orcadian gene pool,
although they noted that it might not be possible to distinguish
2.47 from an Anglo-Saxon contribution in other
parts of Britain. We compared the frequencies found in
the Central English, Frisian, and Norwegian samples of
(1) the 2.47 and 3.65 haplotypes on their own, (2) these
two haplotypes plus their one-step mutational neighbors,
and (3) these two haplotypes plus their one-step networks
(defined as all haplotypes within a sample connected
to the named haplotype by a series of one-step
mutations via observed intermediate haplotypes). As
suggested by the results in the previous paragraph, in
each case the frequency distribution in Central England
more closely matched that in Friesland than that in Norway
.
Thus, neither of these two haplotypes provided any
positive evidence of a (Norwegian) Viking contribution
to the Central English gene pool that could not be ex1018
Weale et al.
plained by a substantial contribution originating in
Friesland only.
Population Genetic Models
We explored various population genetic models
(see Materials and Methods) to evaluate whether or not
a large Anglo-Saxon migration event is needed to explain
the extremely high Central English-Frisian affinity
.
We started with a simple model of population fission
with no background migration. We found a 95% credible
interval for the split date using BATWING of 0–88 generations
(0–2,200 years BP, assuming 25 years per generation),
which corresponded well with the 95% confidence
interval from the Monte Carlo likelihood method
with no background migration (0–91 generations or 0–
2,275 years BP assuming 25 years per generation).
Next, we looked at the levels of background migration,
operating continuously from generation to generation,
needed to maintain the Central English-Frisian
genetic similarity under two other scenarios not involving
Anglo-Saxon mass migration. Under an Island Model
scenario (constant background migration between two
populations that split at TS 5 `) the 95% confidence
interval for m, estimated from the Monte Carlo likelihood
method, is 0.3%–50% (where 50% indicates complete
panmixia and is a maximum value for m). The
same result (to the significant digit given) is found under
a Neolithic mass migration scenario (population split
240 generations BP). We note that a figure of m 5 0.3%
is three times higher than the figure we estimated as
representing an implausibly high value for m, well in
excess of realistic values, based on migration statistics
to and from the European Economic Area as a whole
over the past 25 years. If we set m at the implausibly
high value of 0.1%, the 95% confidence interval for a
Central English-Frisian split date is 0–97 generations
(0–2,425 years BP, assuming 25 years per generation).
The figure of 97 generations BP represents an extreme
upper limit for the migration event in this case both
because it is based on such an implausibly high value
for background migration and because it requires the
most severe mass migration event imaginable, namely a
100% replacement of the English Y chromosome pool.
Next, we assumed that an Anglo-Saxon migration
event did take place 60 generations ago (i.e., 1,500 years
BP assuming 25 years per generation) and asked how
big an event would be needed to explain the Central
English-Frisian genetic similarity. If the Central English
and Frisian populations were very different at the time
of the event, a larger mass migration would be needed.
We therefore started by assuming complete genetic identity
of the two populations at the time of the Neolithic
(i.e., a Central English-Frisian population split 240 generations
BP). Assuming no background migration, the
95% confidence interval of the proportion F of the Central
English population derived from an Anglo-Saxon
mass migration event is 65%–100%. If a background
migration rate since the Neolithic of m 5 0.1% is allowed,
the 95% confidence interval for F widens to
50%–100%. This result is unchanged if a 30-year generation
time is assumed (i.e., an Anglo-Saxon migration
event 50 rather than 60 generations ago).
Discussion
Our results indicate the presence of a strong genetic
barrier between Central England and North Wales and
the virtual absence of a barrier between Central England
and Friesland. Any attempt to explain these results in
terms of demographic history and migration needs to
encompass both these findings satisfactorily. The Central
English-North Welsh barrier cannot be explained
purely as a simple isolation-by-distance phenomenon
because it contrasts strongly with the lack of evidence
for a cline among the five widely separated English
towns. Our findings are particularly striking, given the
high resolution and rapid mutation rate of the Y chromosome
haplotypes on which they are based. These allow
genetic barriers, if they exist, to be clearly defined.
The best explanation for our findings is that the
Anglo-Saxon cultural transition in Central England coincided
with a mass immigration from the continent.
Such an event would simultaneously explain both the
high Central English-Frisian affinity and the low Central
English-North Welsh affinity.
If we use a rate of 0.1%,
as observed over the past 25 years, to represent an extremely
high value for continuous background migration
between Central England and continental Europe, then
we estimate that an Anglo-Saxon immigration event affecting
50%–100% of the Central English male gene
pool at that time is required
. We note, however, that our
data do not allow us to distinguish an event that simply
added to the indigenous Central English male gene pool
from one where indigenous males were displaced elsewhere
or one where indigenous males were reduced in
number. Furthermore, although our models assume a
single instantaneous migration event, we would also expect
a more gradual process lasting several generations
but still resulting in the same degree of admixture (a
picture which may fit the historical data better [Ha¨rke
2002]) to produce very similar genetic patterns.
We accept that our data do not prove conclusively
that an Anglo-Saxon mass migration event took place.
If a background migration rate of 0.3% is allowed between
Central England and Friesland, then the need for
a mass migration event disappears. However, we note
that this is an extremely high rate even by modern standards
and would have to have been maintained continuously
over thousands of years. A background migration
rate of 0.3% would imply that one in six of today’s
Central English males descend from Frisians (or a population
identical to Frisians) that emigrated to England
after the Anglo-Saxon period and that an equal proportion
of today’s Frisians descend from English in a like
manner. We also note that under a unidirectional gene
flow model involving immigration into Central England
only, the rate of background migration would then have
to double to be at least 0.6% on a continuous basis.
It is also true that a mass migration event could
have occurred plained by a substantial contribution originating in
Friesland only.

Anglo-Saxon settlements and culture appeared
throughout England but, importantly, did not extend into
North Wales, where many of the original Celtic Britons
living in England are thought to have fled (Kearney
1989; Davies 1993, 1999). Conflict between the Welsh
and Anglo-Saxon kingdoms continued over a long period.
Offa’s Dyke (an earthwork barrier 240 km long)
was constructed ca. A.D. 790 and provided a well-defined
boundary between England and Wales. The linlinguistic,
cultural, and political separation of the two regions
lasted at least until A.D. 1282 when Edward I of
England defeated the Welsh King Llywelyn II (Davies
1993). Our results suggest that this separation has also
restricted male-mediated gene flow between the two regions
over the past approximately 1500 years.
Comparisons of Central English and Norwegian
haplotypes reveal no evidence of distinctive common
signature haplotypes indicative of Viking origin, in contrast
to Orcadian-Norwegian comparisons (Wilson et al.
2001). However, the Vikings who may have settled in
East Anglia and the Midlands are thought to have been
predominantly from Denmark, rather than Norway
(Richards 2000). Previously published data suggest that
the Danish have greater Y chromosome genetic affinity
with the English than with the Norwegians (Malaspina
et al. 2000; Rosser et al. 2000). However, the Danish-
German border is believed to be another source location
of the Anglo-Saxons (Kearney 1989; Davies 1999), so
any Danish Viking influence on the English gene pool
may prove difficult to distinguish from Anglo-Saxon influence.
Further studies within Scandinavia and elsewhere
are needed to resolve this issue.
This study shows that the Welsh border was more
of a genetic barrier to Anglo-Saxon Y chromosome gene
flow than the North Sea. Remarkably, we find that the
resultant genetic differentiation is still discernible in the
present day. These results indicate that a political boundary
can be more important than a geophysical one in
population genetic structuring and that informative patterns
of genetic differentiation can be produced by migration
events occurring within historical times.


http://class.csueastbay.edu/anthropologymuseum/2006IA/DNA_PDFS/yDNA/Weale2002.pdf


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Postby yialousa1971 » Sat Mar 26, 2011 9:49 pm

Conclusion : samples from central England and Friesland are almost undistinguishable. proving that a massive proportion of central English people descend from the Anglo-Saxons. Conversely, samples in North Wales and Norway were quite different.


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Postby yialousa1971 » Sun Mar 27, 2011 7:42 pm

Wheres dog gone, run off I think.
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Postby supporttheunderdog » Sun Mar 27, 2011 10:29 pm

No not running at all: I have a life outside of CF and, having been away from home for nearly a week, was living it by spending some quality time with my ladies, all of whom were in need of some TLC.

1) re you 21,15 post on vikings can you give me a URL or other reference to where I can find the study?
2) Re the UCL 2002 Study I am aware of it and had read what you had posted before I had started this thread.
That Study has been superseded by at least two other later studies which I referred to in my opening item - I am in course of putting togther a more comprehensive response with links to supporting items /these latest studies and will revert but it might take a few days.
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Postby yialousa1971 » Sun Mar 27, 2011 10:45 pm

supporttheunderdog wrote:No not running at all: I have a life outside of CF and, having been away from home for nearly a week, was living it by spending some quality time with my ladies, all of whom were in need of some TLC.

1) re you 21,15 post on vikings can you give me a URL or other reference to where I can find the study?
2) Re the UCL 2002 Study I am aware of it and had read what you had posted before I had started this thread.
That Study has been superseded by at least two other later studies which I referred to in my opening item - I am in course of putting togther a more comprehensive response with links to supporting items /these latest studies and will revert but it might take a few days.


http://mbe.oxfordjournals.org/content/25/2/301.full

LOL you're putting togther a more comprehensive response. You need your eyes testing if you can't see the population of Britain has multiple origins!
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Postby supporttheunderdog » Sun Mar 27, 2011 11:14 pm

yialousa1971 wrote:
supporttheunderdog wrote:No not running at all: I have a life outside of CF and, having been away from home for nearly a week, was living it by spending some quality time with my ladies, all of whom were in need of some TLC.

1) re you 21,15 post on vikings can you give me a URL or other reference to where I can find the study?
2) Re the UCL 2002 Study I am aware of it and had read what you had posted before I had started this thread.
That Study has been superseded by at least two other later studies which I referred to in my opening item - I am in course of putting togther a more comprehensive response with links to supporting items /these latest studies and will revert but it might take a few days.


http://mbe.oxfordjournals.org/content/25/2/301.full

LOL you're putting togther a more comprehensive response. You need your eyes testing if you can't see the population of Britain has multiple origins!


Thanks - I'll look it out.

The recent studies I am am aware of and referring to, Sykes and Oppenheimer in particular, themselves recognise the diverse genetic input in Britain:one can even trace the Balkans origin of some genetic lines, probably introduced with Roman Auxiliary troops/Mercenaries, and there are some suggestions of African input in one place in Yorkshire, possibly again related to slaves, etc, who accompanied Roman Legions.
Indeed one study suggests that about a 1000 years ago England was genetically more diverse than it is today.

Its the same in Greece- almost certainly multiple waves. That however is another topic.

These studies also recognise that the genetic make up can vary quite considerably from place to place. One reason you might for example find a predominance of Vikings in the Lake district or the uplands in Yorkshire is that it is hard country to farm and it was all that was available to them.

Same may well go for Orkney's' and Shetlands' -

What the debate is about is the current extent of that diversity, where the most recent studies by Sykes, Oppenheimer and all, suggest that Anglo-Saxon Origin theory is grossly overstated, and that that majority of the English/British alive today are descended principally from the peoples who came to Britain along the Atlantic coast from Iberia around the end of the last glacial Maximum.

More to follow.
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Postby Piratis » Sun Mar 27, 2011 11:43 pm

supporttheunderdog, are you English?
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