Genetic Diversity
in Flemish Y-DNA
Guido
Deboeck
Abstract
More than a
million Americans are descendants of Flemish immigrants. Close to 400,000 Flemish immigrants settled
between Michigan and Oregon in the late 19th and early 20th centuries. Flemish people also had
major influences on the English, the Scots, and the Irish.
Genetic
genealogy is a relative young field that has made interesting discoveries
regarding surname relationships. Surname
relationships are, however, dependant on the origin of the surname, changes in
spelling of surnames, interpretations and adaptations of surnames to new
environments. In contrast, geographical DNA projects focus on a particular
geographical area where people have experienced a common history over
time. The interweaving of history,
conventional, and genetic genealogy often produces a better picture of the
evolution of a group of people. A first
step in this direction is to discover the main haplogroups present in a
particular population. This article
focuses on refining the understanding of the distribution of haplogroups among
people with Flemish roots, creating a baseline for the study of the mosaic of
genetic diversity in Europe.
Address for correspondence: Guido
Deboeck, gdeboeck@mac.com
Received: February
26, 2008; Accepted: March 23, 2008.
Introduction
The
ancestors of the Flemish originally came from Flanders, a region located in what is
today the northern part of Belgium.
Flanders has an area of 5,221 square miles
(13,522 square km), roughly the size of Connecticut.
The current population of Flanders is slightly over 6 million, resulting in a population density
of 886 per square mile (342 per square km).
The
history of Flanders and of the Flemish people in
particular provides an interesting background for the study of Flemish DNA that will lead to better
understanding of the mosaic of European DNA.
By interweaving history, genealogy, and genetic genealogy, part of this
mosaic of Flemish DNA can be revealed.
This
paper starts with a brief historical overview of Flanders and the Flemish people, and
subsequently describes the datasets of Y-chromosome STR haplotypes assembled for the
present study. Finally it provides the
frequency distribution of the major haplogroups and discusses how this is
different from previously published papers.
The paper concludes by bringing together the origin of the main
Y-chromosome haplogroups found in the Flemish, and discusses their European
context. Portions of this article were
previously published in "Flemish DNA & Ancestry" (Deboeck,
2007).
Historical Background
Centuries
ago the deltas formed by three great rivers, the Weser, the Meuse, and the Scheldt, were inhabited by the Goths,
who were an ancient Teutonic people.
These rivers deposited their muddy debris for ages on what became known
as “the Low
Countries.” The fertility of these plains became fatal to
the native possessors because many foreign tyrants sought to conquer these Low Countries.
This aroused and inspired resistance among the inhabitants; they
acquired the genius of liberty and of upholding independence. These natives, whose name became synonymous
with liberty, were cultivators of
soil; the tribe was called Frisii, Frisories or Frisians.
Some four hundred years after the
fall of the Roman
Empire,
Charles the Bald, the first king of France, established the County of Flanders as a feudal fief. For five centuries Flanders remained independent of foreign
policies and decisions. Thereafter, from
the 13th till the early part of the 19th century, the southern part of the Low Countries was occupied by foreign
forces. Arnold Flemming
wrote: “Happy is the land that has no history; no history,
that is, of blood and strife.
Such cannot be said of Flanders, it is full of history” (Flemming,
1930). The history of Flanders is a sequence of periods (see Table 1) in which Flanders was dominated by the Spanish, the
Austrians, the French, and the Dutch, before it became a region or state of Belgium.
Table 1 Historical Overview of Flanders’ Struggle for Independence
|
|
|
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Burgundian Netherlands
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1384-1482
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Habsburg Netherlands
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|
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Spanish Netherlands
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1556-1713
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|
Austrian Netherlands
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1713-1789
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|
United States of Belgium
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1790
|
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Austrian Netherlands
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1790-1794
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French Republic
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1795-1804
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French Empire
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1804-1814
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United Kingdom of Netherlands
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1815-1830
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Kingdom of Belgium (unitary state)
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1830-1993
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Flanders: federal region/community
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1993 - present
|
Figure 1 Map of the southern Low Countries in the late Middle Ages, including present-day Flanders (Source: Carson, 1990)
In 1830 the big powers of Europe created Belgium as an artificial state
(other artificial states, which have fallen apart since their creation, were Yugoslavia and Czechoslovakia).
Belgium was originally set up as a
unitary state with a parliamentary system of government. Through four revisions of the constitution,
the unitary state was transformed into an ad hoc federal state, with
three regions responsible for territory-related matters and three communities
responsible for personal matters. The
complex socio-economic and political structures of the Belgium Federal Model
remain an “unfinished design.”
Flanders now refers to that part of Belgium that is inhabited by speakers of
Dutch or Flemish dialects, and the inhabitants are considered to be
Flemish. A broader definition of “the
Flemish” includes people who have Flemish roots, but have migrated to Britain, Canada, the U.S., or elsewhere. This broader definition was adopted in this
study so that DNA samples collected from people living in the U.S. and elsewhere could be used and
compared to DNA derived from sources in Flanders.
Materials and Methods
This
study relies on the pooling of three sets of Y-chromosome STR data. The two larger datasets contain data on 12 Y-STR markers, DYS393, DYS390, DYS019, DYS391, DYS385a, DYS385b, DYS439, DYS389i, DYS392, DYS389ii, DYS437, and DYS438. The Flanders-Flemish DNA (FFDNA) project dataset has
haplotypes with 12, 25, or 37 markers from Family Tree DNA’s (FTDNA’s)
three panels of markers. Each set was
collected in a different way from different subjects and is independent of the
others.
The first
set of Y-STR haplotypes was derived from the FFDNA project, launched by the author
in November 2005. As of March 2007, 45
records had been submitted to this project at FTDNA. These records were mainly from people living
in the U.S. who believed that their
patrilineal lines came from Flanders. These initial 45
records were used for the first analysis undertaken in the spring of 2007 and
published in September 2007 (Deboeck, 2007).
Since September 2007, 18 more records have been added to the FFDNA
project, bringing the total number of FFDNA samples to 63. While all of the participants who have joined
the FFDNA project believe that their paternal lines come from Flanders, there is no way to verify this.
The map in Figure 2 is marked to indicate where members of the FFDNA project
live. The map in Figure 3 shows the location of the most distant ancestors of those
members. On the second map, three
markers are shown in Ireland, but these are from people who
believe that their ancestors migrated there from Flanders.
Arnold Fleming wrote that “history is an eternal tangle of cross
purposes, and we could not take a more complicated case than the heterogeneous
factors lying behind the influence of the Flemings upon the British people”
(Fleming, 1930).
Figure 2 Residences of Flanders-Flemish DNA Project members.
Figure 3 Residences of the earliest known ancestors of
Flanders-Flemish DNA Project members.
In addition to the influence on Britain, Flemish emigration in the 19th
and early 20th centuries has resulted in over one million people of Flemish
descent currently living in the U.S. (Denys,
1984; Baye, 1987).
These Flemish Americans clearly have Flemish ancestors (van Molle L., Pansaerts C,
1996). In sum, the maps and A. Fleming’s
work show that in the U.S., England, Scotland, and Ireland there may be a lot of people who
have ancestors who originally came from Flanders.
The second set of haplotypes is
based on 112 DNA samples extracted from blood samples taken at the university hospital
of the Catholic University of Leuven (KUL). According to Professor Ronny DeCorte, these blood samples were mostly taken from people
who were from Leuven. Each of
these 112 haplotypes contains 12 markers.
The modal values of the 12 markers
for these 112 haplotypes correspond exactly with the modal values found in the
45 records collected via the FFDNA project.
The fact that the modal values correspond so closely provides evidence
that the data collected via the FFDNA project indeed represents DNA from people with Flemish roots.
The third data set was derived
from a genetic study of 113 unrelated Flemish males, who were typed for 12
Y-STRs. This dataset was obtained from
G. Mertens and appeared in the Fall 2007 issue of this
journal. The main result of that study
showed that in this sample of 113 Flemings, there were 56.6% belonging to
Haplogroup R1b, 19.5% belonging to Haplogroup I1 (I-M253), 8% to Haplogroup
I2b2 (I-M223), 4.4% to Haplogroup E1b1b1 (E-M35, formerly E3b), and the
remainder to Haplogroups G2a (G-M201), I2a (I-P37), J2a1b (J-M67), J2a1k (J-DYS445<7), J2b (J-M12) and L,
ranging from 0.9 to 3.5% of the total (Mertens, 2007).
Merging
all three data sets provided a total of 288 haplotypes with complete data for
the ten markers DYS393, DYS390, DYS19, DYS391, DYS385a&b, DYS 439, DYS389-1&-2, and DYS392. This combined Flemish Y-STR dataset was then used to update
the analyses previously carried out by Deboeck (2007) and Mertens (2007).
The
methodology that was used consisted of the following steps:
1. All
three data sets were merged into a single spreadsheet. The identifier and source of each haplotype
was kept in a separate column.
2. The
Haplogroup Predictor program (Athey, 2005; Athey, 2006) was used in batch mode
to predict the haplogroups for the KUL-Leuven data
set, and to reanalyze the haplogroup for the other two datasets so that all of
the predictions would have the same basis.
3. The
data records were sorted by source and haplogroup to compute the haplogroup
frequencies for each data set.
4. The
overall haplogroup frequencies were calculated for the combined dataset.
5. The
allele frequencies were calculated for each marker in the combined dataset for
Haplogroups R1b and I1.
6. The
modal haplotypes for the most prevalent haplogroups in the combined Flemish
dataset were determined.
7. For
each of the most prevalent haplogroups, the difference in origin and history of
the haplogroups were compared.
Results
The main
results from applying the above steps produced a table of allele frequencies
for 37 markers in 166 Flemish R1b haplotypes and 45 I1 haplotypes as shown in Tables 2 and 3; Table 4 shows the haplogroup distributions from the three datasets,
both separately and combined; a series of pie charts in Figures 4-7 show the distribution of haplogroups in these datasets;
and Table 5 compares the Flemish
modal haplotypes with the modal haplotypes of the most common haplogroups.
Table 2 shows the allele frequency distributions for Flemish
members of Haplogroup R1b, while Table 3
presents similar data from Haplogroup I1.
However, note that for some markers there is insufficient data for an
accurate calculation of the allele frequencies or even the modal value—these
are subject to change as more data is accumulated. Table
4 provides the haplogroup distributions for the pooled Flemish haplotypes.
A
comparison of Figures 4 and 5 shows
that the data obtained via the FFDNA project corresponds closely to the 113
records used by G. Mertens in a study of paternity cases in Belgium. The dominant haplogroup in both cases was
R1b, which represents between 54% and 56.6% of the total. The second largest haplogroup in these two
samples is Haplogroup I1 which represents between 19.5% and 25% of the
total. Other haplogroups that occur are
E1b1b, representing between 2% and 4.4%, and Haplogroup G2a, representing
between 3.5% and 7% of the population. A
difference between the FFDNA sample and the Mertens dataset is in the
occurrence of Haplogroup R1a, which represents 7% in the first sample and does
not appear in the second, though if the true frequency is only 1-2%, this is
not a surprising result.
Figure 6 provides the detailed
distribution of haplogroups found among the 289 Flemish haplotypes. The haplotypes from all three data sets were
submitted to the 2007 21-haplogroup version of Whit Athey’s Haplogroup Predictor program
(Athey, 2005, 2006) so that all of the predictions would have the same
basis. This produced a number of changes
from their old designations, some of which simply involve newer nomenclature
(ISOGG, 2008); these changes are reflected in Table 4, which was used to produce Figure 6.
Combining
the subhaplogroups' frequencies into the major haplogroups produces the
distribution shown in Figure 7 and
allows direct comparison with the G. Mertens results. An analysis of the full set of 288 haplotypes
shows that 57.6% are Haplogroup R1b, 15.3% are I1 (old I1a), 5.2% are J2, 4.5%
are E1b1b (old E3b), 4.2% are R1a, and 3.5% are G.
Table 5 compares the Flemish (R1b) Modal
Haplotype (FMH) with the modal haplotypes of other R1b groups. Where the allele value of a particular marker
in the FMH differs from the allele value of the other modal haplotypes, the
values are shown in red font. This
comparison shows that the FMH differs on only three markers from the Atlantic
Modal Haplotype (AMH). On DYS449 the FMH has a value of 30
while the AMH has 29; on DYS456 the FHM has a value of 16 while the AMH has 15, and on CDYa the FMH has a value of 36 while the AMH has 37. It should be noted that for CDYa, there is very little difference in the FMH allele
frequency for a value of 36 versus 37.
A
comparison of the FMH with the modal haplotype of the Frisian R1b cluster of
McEwan (2006) shows differences on the markers DYS390, DYS447, DYS449, GATA-H4, DYS456, DYS576, and CDYa/b,
plus differences in the complex marker, DYS464.
Other
comparisons: a comparison of the FMH with the R1b model values from the German
geographic project at FTDNA shows that they differ on only one marker, DYS449. A comparison of the FMH with the R1b modal
values from the Spanish geographic project at FTDNA shows that they differ on
five markers, DYS439, DYS458, DYS464d, GATA-H4, and DYS456. A comparison with the R1b modal from the British Isles geographic project at FTDNA shows
that the FMH differs on three markers with the British Isles R1b modal (DYS 449, CDYa,
and CDYb. In
summary there are some differences between the FMH and other major R1b
population groups in Europe, which invites further investigation.
Table 6 shows a similar comparison for
the Flemish I1 modal haplotype. The
modal values for many of the markers are based on fewer than 10 haplotypes, so
no firm conclusions may be drawn for these markers. However, one notable difference with the
Frisian I1 modal haplotype (McEwan, 2006) is on DYS390, where adequate data
exist. The Flemish I1 modal haplotype
has a value of 24 on this marker, matching the overall European I1 modal
haplotype in this case, while the Frisian I1 modal haplotype has a value of 23.
Discussion
What are
the main differences between the haplogroups that are most prevalent among the
Flemish?
Spencer
Wells’ book, Deep Ancestry, describes
the haplogroups as follows:
• Haplogroup R1b members are descendants
of someone who 30,000 years ago gave rise to the defining marker M343. These descendants that dominated the human
expansion into Europe are likely the same as the Cro-Magnons
(Wells, 2007, p 226).
• Members of Haplogroup I1 have the
defining genetic marker M253, and moved 15,000 years ago from their refugia in the Balkans to the north of Europe.
This haplogroup is still very common in Scandinavia and hence it is likely that many
Vikings descended from this line. The
Viking raids on the Low Countries and Britain help to explain the dispersal of
this lineage (Wells, 2007, p 213-14).
• Haplogroup E1b1b, characterized by the
M35 marker, appeared in the Middle East among populations of the first farmers who spread
agriculture from the Middle East to Europe (Wells, p209).
• Haplogroup G2a (referred to as G2 in the
book), identified by the marker P15, arose in he Middle East and spread westwards through
modern Turkey into southeast Europe.
Most migrations took place more than 15,000 years ago before much of Europe was locked in ice during the last
glacial maximum. When the glaciers
finally began to recede, the G2a lineage expanded northward and eastward to
repopulate Europe (Wells, 2007, p 211).
It should
be noted that almost all haplotypes classed as R1b in this geographic study are
likely also to be R1b1b2-M269. Most of
the present-day European males with the M343 marker also have the P25 and M269
markers. These markers define the R1b1b2
subclade (ISOGG, 2008). The subclade
R1b1b2g (defined by S21/U106), although recently discovered, appears to be the
most common downstream marker from R1b1b2, appearing in over 35% of those
R1b1b2 people tested. This group has a
maximum in Frisia (the Netherlands) and, in general, is the
predominant R1b sub-haplogroup in northwest Europe.
It is also found in Belgium, in the UK, in southern Sweden, Switzerland, and Italy.
It only sporadically appears in Spain, Poland and Lithuania (Polish), Finland (Swedish) and Romania (Romanian). The S21/U106 subclade may have originated
near the end of the last ice age as Europe began to be repopulated, or perhaps more or less
9000 years before the present, possibly in the northern European mainland.
In
summary, the analysis of three datasets of Flemish Y-STR values, collected from three
independent sources, confirm the dominant presence of Haplogroup R1b in Flanders.
In addition, Haplogroup I1-M253, and to a smaller extent Haplogroups
I2b1-M223, J2, E1b1b, R1a, G2a, and I2a, have been shown to occur among Flemish
people. The data collected via the FFDNA
project, which was mainly submitted by people living in the USA, is fairly representative of the
overall distribution of Flemish DNA.
Hence there is continuity in the paternal lines of the Flemish and
Flemish-Americans. The genetic
characteristics of a small population living in the north of Europe provide a glimpse into the
complexity of the genetic mosaic in Europe.
Future
research should concentrate on drawing more in-depth comparisons of Flemish DNA with the patterns observed in
Frisian, German, British, Spanish and French populations. Based on the history of Flanders it is
reasonable to assume that the German influence on Flemish DNA is likely to be much stronger
than, say, the French. The British
influence on Flemish DNA is likely to be minimal since Britain was never an occupying power in Flanders.
However Flemish DNA may be found back on the British Isles since centuries ago there were
large migrations of Flemish people to England, Scotland, and Ireland.
This should stimulate genetic genealogists interested in the British Isles to trace their ancestry to the
continent, in particular to Flanders, that small piece of land across the Channel that has
been a battleground for centuries and continues to be the center of attention
as to its influence on the political unity of Europe.
Acknowledgements
The
author is grateful for the advice and collaboration he received from Whit Athey, Gerhard Mertens, Paul Duran, and Prof. Ronny DeCorte.
Web
Resources
http://www.hprg.com/hapest5/
Haplogroup Predictor Program
http://www.familytreedna.com/public/Flanders/
Flanders-Flemish DNA Project
web site
http://www.senate.be/english/federal_parliament_en.html
Belgian Federal Model
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