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Tuesday, April 27, 2010

The Anthropology of Baltic-Finns(2)

The Origin of the Baltic-Finns from the
Physical Anthropological Point of View
Markku Niskanen1
University of Oulu, Finland
ΙΙ
Genetic Relationships in Light of
Nuclear Gene-Frequency Evidence
I used the Chi-Square distance measure of the SPSS-program
to calculate genetic distances between populations from the
nuclear gene-frequencies. The raw data is collected from
literature (Nevanlinna 1973, Kajanoja 1978, Zubow 1979,
Eriksson 1988, Heapost 1993, Cavalli-Sforza et al. 1994). The use
of this distance measure does not tolerate missing values. For
this reason, I had to vary the markers used depending on which
populations were compared and, therefore, the distance values
computed using different sets of markers are not comparable. In
general, the larger the number of genetic markers used, the
more reliable the genetic distance.
The Finns and the Estonians represent the Baltic-Finns in
these analyses. The Finns represent the Baltic-Finns when large
numbers of genetic markers are needed. I had to pool the Mari
and the Komi to represent the Finno-Ugrians of the easternmost
Pasted Graphic 3.pict
Europe (EFU = Eastern Finno-Ugrians) due to insufficient genefrequency
data.
At first, I calculated the genetic Chi-Square distances of
European populations from the Finns, Saami, the European
average, inhabitants of modern Turkey (Anatolia), Mongols,
and Japanese to determine how these populations are placed
within the genetic landscape of Eurasia (Table 5). These
distances reveal that the Finns are genetically close to their
Germanic-speaking neighbors (the Swedes, Germans, and
Norwegians), the least distant population from the Saami, and
no more distant from the European average than are the Irish
and the Basque. Based on the FST genetic distances presented by
Cavalli-Sforza et al. (1994:270), the Finns are genetically a little
closer to the Belgians (FST = 63), Germans (FST = 77), and
Austrians (FST = 77) than to the Swedes (FST = 82). This
difference is a result of Cavalli-Sforza et al. (1994) using a
different genetic distance measure and computing distances
using a data set including missing values. As Nei and
Roychoudhury (1993) have pointed out, inter-group distances
can be misleading if the data set from where the distances are
computed includes missing values.
The genetic distances to the inhabitants of modern Turkey
(Anatolia) indicate that the Finns are just like other North
Europeans genetically more distant from the Near Eastern
people than are the Central-and South Europeans. The genetic
distances to the Mongols and the Japanese indicate that the
Finns are genetically somewhat less distant to these eastern
populations than the Europeans are as an average, but not any
closer than other Europeans living along the same longitude
(for example, the Greeks and Bulgarians in respect to their
distances to the Japanese). Many allele frequencies of the Finns
and especially the Saami, which differ from the European
averages, and the surprisingly small genetic distance off the
Icelanders from the Mongols (which is smaller than that
between the Finns and the Mongols) are most likely results of
random genetic drift in partially isolated and numerically small
population.
The FST distances (above the diagonal) and Chi-Square distances
(below the diagonal) were used to examine the genetic
relationships of the Finns with other North European
populations more closely (Table 6). These two genetic distance
Pasted Graphic 4.pict
measures provide very similar pictures of the inter-population
genetic relationships. Based on these distances, the Swedes are
the least distant from the Finns. The FST and Chi-Square
distances between the Finns and the Swedes (82 and 156) are,
however, considerably larger than the distances of the Swedes to
the Norwegians (18 and 84), Danes (36 and 98), Dutch (41 and
118), and English (37 and 123), but about equal to those of the
Swedes to the Irish (94 and 192) and Scottish (74 and 157). This
finding indicates either that the Baltic Sea has formed a partial
restriction of gene flow and, therefore, a genetic boundary, or
that the Northwest Europeans and the Northeast Europeans
have partly different genetic origins. The relatively large
geneticdistances between the Finns and other Northwest
Europeans could also have resulted from genetic drift, which
has altered the Finns’ allele frequencies.
The genetic distances of the Finns to the Saami are smaller
than those between the Saami and other European, but still very
large. This finding indicates that the Finns and the Saami were
originally more distant from each other, but have exchanged
genes during the more recent history. In other words, it appears
that these two Finno-Ugrian-speaking people most likely do not
descend from a common ancestral population which lived only
4,000-5,000 years ago. The same conclusion has also been
reached in light of the mitochondrial DNA evidence discussed
below.
I computed genetic Chi-Square distances between the Irish,
Swedes, Finns, Saami, pooled Mari and Komi (EFU = Eastern
Finno-Ugrians), Samoyeds (Nenets and Nganasans), and
Mongols to estimate the genetic relationships between the
Uralic-speaking populations and their non-Uralic-speaking
neighbors. Based on these distances (Table 7), the Finns
representing the Baltic-Finns are genetically closer to both the
Swedes (99) and the Irish (136) than to the pooled Mari and
Komi (142). Interestingly, the genetic distance between the
Finns and the Irish (136) is not much smaller than the distance
between the Irish and the pooled Mari and Komi (159)
regardless of the geographic distances between the groups in
question.
The genetic distances in Table 7 also reveal that the Finns,
Saami, and the pooled Mari and Komi (EFU) to the Samoyeds
(Nenets and Nganasans) are not much smaller than those of the
Pasted Graphic 5.pict
Pasted Graphic 6.pict
Pasted Graphic 7.pict
Irish and the Swedes to the Samoyeds. The Samoyeds are also
genetically equally distant from the Europeans (both Uralic- and
non-Uralic speakers) than are the Mongols. These findings
demonstrate clearly European gene pools of even the
easternmost Finno-Ugrians and North Asian gene pools of the
Samoyeds. It is clear that the ancestors of the Finno-Ugrians and
the Samoyeds either diverged a very long time ago from each
other (tens of thousands of years ago) or that there never was a
common Uralic homeland. A study of the Samoyeds’ Ychromosomal
DNA variation by Karafet et al. (1997) supports
the idea of distinct genetic and geographic origins for the
Samoyeds and other Uralic-speaking populations.
I had to reduce the number of genetic markers used to
include the Estonians in the genetic distance analyses. Results
presented in Table 8 are, however, similar to those presented
above. The Baltic-Finns (the Estonians and Finns) are
genetically somewhat closer to the Swedes (144 and 116) than to
the pooled Mari and Komi (146 and 149). This finding indicates
that the Baltic-Finns and the more eastern Finno-Ugrians do not
have recent common ancestors who lived only about 3,000 years
ago between the Volga River and the Ural Mountains, as argued
in the traditional migration theory. If these groups had a
common homeland, they must have diverged from each other
over ten thousand years ago instead of 3,000 years ago. The
genetic distances presented also demonstrate that the Finns are
genetically closer to the Swedes (116) than to the Estonians
(118), indicating that the Finns and the Estonians probably
diverged from each other (through the migration of the Finns’
ancestors from Estonia) earlier than the traditionally assumed
migration date (during the first millennium AD).
The Mitochondrial DNA
and Y-Chromosomal DNA Evidence
The mitochondrial DNA (mtDNA) is inherited matrilineally,
whereas the Y-chromosomal DNA is inherited patrilineally. This
lack of genetic recombination allows reconstructions of
maternal lineages (from the mtDNA data) and the paternal
lineages (from the Y-chromosomal data) from the DNA
sequence differences. This data also provides information about
more ancient population history than the nuclear gene
frequencies (of blood groups and enzyme polymorphism),
which can change markedly through genetic bottlenecks
(genetic drift).
The mtDNA studies (Sajantila et al. 1995, Lahermo et al.
1996) reveal that the non-Saami Finno-Ugrians of Europe (the
Finns, Karelians, Estonians, Volga-Finns) have the same genetic
origin as the non-Uralic-speaking Europeans, and that the Saami
represent a unique and ancient sub-group of Europeans that
had separated from the other Europeans over 10,000 years ago.
Therefore, the Baltic-Finns (the Finns, Karelians, and Estonians)
and the Saami do not appear to descend from a common
ancestral population that lived as recently as a few thousand
years ago. The genetic admixture between the Baltic-Finns and
the Saami is also rather recent, but adequate enough to make
the Finns and the Karelians the closest genetic relatives of the
Saami.
Studies of the mtDNA and Y chromosomal DNA reveal
information about ancient population movements and
directions. For instance, the distribution and diversity of the
mtDNA haplogroup V indicates a population expansion from
Southwest Europe starting about 15,000 years ago over the
western half of Central Europe and the entire Northwest
Europe. The existence of this haplogroup among the Saami,
Baltic-Finns, and the Volga-Finns indicates a sizeable ancient
West European genetic component in their gene pools. Its
frequency is the highest among the Saami (40.9%), Basque
(20.0%), and Catalonians (26.7%). It exhibits the greatest
amount of diversity among the Iberian populations, suggesting
that this region is its most likely area of origins. This diversity is
far less among the Saami, indicating that their extremely high
haplogroup V frequencies resulted from founders’ effect
(Torroni et al. 1998).
While studies of the mtDNA variation indicate strongly the
western maternal gene pools of Europe’s Finno-Ugrians, the
studies of Y-chromosomal DNA variation indicate eastern
paternal genetic contribution in their gene pools in addition to
the western one. Kittles et al. (1998) noted significant
differences in Y chomosome haplotype variation between
western and eastern Finland. They thought that the western
Finns, characterized by the haplotype B, were of predominantly
western ancestry and that the eastern Finns, characterized by the
haplotype A, were of Asian ancestry because the haplotype A
contains the DYF155S2 deletion found most commonly among
Asians. According to Jobling et al. (1996), this deletion is,
however, also encountered among the Norwegians and Greeks,
and it is the most common in Finland and Mongolia. Therefore,
I argue this deletion could have emerged anywhere between
Scandinavia and Mongolia, making it a possible genetic marker
of a late Ice Age population expansion from the South Russian
Plain from where the Northeast Europe and Northwest Asia
were colonized after the Last Glacial Maximum.
Zerjal et al. (1997) argued that the C allele on the Y
chromosome of some North Europeans and North Asians
emerged in Asia and spread to North Europe, where its
existence indicates a considerable paternal genetic contribution
of North Asians to North European populations, especially the
Finno-Ugrians. Villems et al. (1998) discovered that this allele
exhibits more variation among Europeans than among Asians
and, therefore, they suggested that it originally emerged among
Proto-Finno-Ugric populations of Eastern Europe from where it
spread to some Siberian populations. I consider this C allele as
the genetic marker of the late Ice Age (after the Last Glacial
Maximum) population expansion from the South Russian Plain.
A study of Y-chromosomal markers indicate that the
Samoyeds are descendants of Paleoasiatic populations which
were (linguistically) assimilated by populations who came from
southern Siberia (Karafet et al. 1997). This finding explains the
phenotypic and genetic distinctness of the Samoyeds from other
Uralic-speaking people and similarity with more eastern people,
as well as contradicts the assumption that the Samoyeds are
genetically the closest to the ancestral Uralic people.
All of this craniometric, nuclear DNA, mtDNA, and Ychromosomal
DNA information indicates that the Finno-
Ugrians of Europe originated in Europe, the Baltic-Finns are
genetically closer to the Scandinavians than to the Finno-Ugrian
people in the east, the Saami form a distinct subset within the
Europeans, and the Samoyeds have a distinct genetic origin
from other Uralic people. I will next discuss these findings in
light of the archeological evidence of population history.
The Genetic Origins
and the Archaeological Evidence
Genetic relationships are results of a long history of
population events. These events cannot be reconstructed
accurately without information about the prehistory and history
of the populations in question. In case of northern Eurasia, we
have to reconstruct the population events at least since the Last
Glacial Maximum (23,000-19,500 BC), after which the entire
northern Eurasia was colonized. North Europe was colonized
primarily from two of the Last Glacial Maximum refugia areas:
the Franco-Cantabria in the west and the South Russian Plain
(mainly the Dnieper-Don region). These two refugia areas were
separated from each other by a most likely uninhabited isthmus
located between the Fennoscandian and Alpine glaciers,
although there were some settlements in eastern Austria and
western Hungary. West Siberia and Central Asia (except
southern Urals) were either uninhabited or sparsely inhabited
because West Siberia was almost entirely covered by an
enormous inland lake, and the region between the Caspian Sea
and the mountains of Central Asia was a dry and inhospitable
desert. The upper Yenisey and the Lake Baikal regions were,
however, inhabited, as was the entire southeastern half of
Siberia (Soffer 1990).
Human settlement started to spread over northern Eurasia
when the climate started to ameliorate around 19,500 BC.
Groups originating from Franco-Cantabria settled the British
Isles and the North European Plain located west of the Vistula
River (Creswellian/Hamburg cultures) and the Central
European hill region between the Atlantic Ocean and the
Carpathian Mountains (Magdalenian culture). Groups
originating from the Dnieper-Don region (for example,
Swiderian culture) settled the entire northeastern Europe
located east of the Oder River and possibly westernmost Siberia
(Dolukhanov 1986, 1993, Soffer 1990, Schild 1996). The mtDNA
haplogroup V is the genetic marker of the colonization of
Northwest Europe from Franco-Cantabria (Torroni et al. 1998)
and the Y-chromosomal C allele is the genetic marker of the
colonization of Northeast Europe and West Siberia from the
South Russian Plain (Zerjal et al. 1997, Villems et al. 1998).
The Dnieper-Don region of the South Russian Plain is the
only candidate for a region from where the entire distribution
area of the Uralic languages was settled. The two competing
traditional Uralic homelands – the northeastern corner of
Europe between the Volga Bend and the Urals and Northwest
Siberia – received their first permanent human settlement from
this region. Northwest Siberia also received settlers from further
east of Siberia. It is, therefore, possible that the late Upper
Paleolithic period inhabitants of the entire Northeast Europe
and West Siberia spoke ancestral dialects of the Uralic languages
(Nu–ez 1987, Julku 1995). These groups could have maintained
their linguistic affinities through cultural contacts (marriage
exogamy, etc.) facilitated by the extensive river network of this
huge territory (see Fig. 7 in Nu–ez 1987:14).
There is one problem with the idea that the South Russian
Plain was the Uralic homeland. According to Mallory (1989), for
example, it was possibly settled by Proto-Indo-Europeans already
during the Mesolithic period. A solution to this problem could
be the late Ice-Age northward displacement of ecological zones
and human populations adapted to these zones. The individual
group territories became at first stretched in a north-south
direction, making the seasonal migrations between the northern
and southern parts of hunting territories gradually too long,
which resulted in abandonment of the southernmost territories.
Groups speaking ancestral dialects to Proto-Indo-Iranian could
have expanded into territories being gradually vacated by
speakers of Proto-Uralic dialects through the same process,
either from the northeastern Balkans or the Kuban region east
of the Black Sea.
The earliest post-Last Glacial Maximum inhabitants of the
southeastern shores of the Baltic Sea belonged to the Late
Upper Paleolithic Swiderian culture (10,800-9700 BP / 9800-
8700 cal BC), which descended from the East Gravettian
cultures of the South Russian Plain. This culture spread to the
Oder River in the west (Schild 1996). If people of the Swiderian
culture spoke a Proto-Uralic language, the Oder River formed
the westernmost boundary of the Uralic language at the end of
the Paleolithic period.
Whether or not the people of the Swiderian culture were
Proto-Uralic-speakers, the most ancient human settlement of the
region historically inhabited by the Baltic-Finns belonged to the
Early Mesolithic Kunda culture (emerged about 8700 BC), the
roots of which are in the Swiderian culture. Finland received her
first permanent settlers from the Kunda culture’s territory in the
south (Estonia) and east (Karelia) about 8500 BC. When these
possibly Proto-Uralic-speaking groups arrived in northern
Finland around 8000 BC, they met coastal groups, who had
arrived in northern Fennoscandia as early as 9000 BC from then
ice-free region between the British Isles and Denmark by
following the ice-free coast of Norway. At that time, the inland
regions of central and northern Scandinavia were still covered
by ice (Nu–ez 1987, this volume). The coastal people may have
become Uralic-speaking Saami as a result of cultural contacts
with Proto-Uralic-speaking populations inhabiting inland
regions of northern Fennoscandia. This language shift would
explain why the Saami are genetically so distinct from
Scandinavians and Baltic-Finns, but speak languages that are
quite similar to the Baltic-Finnish languages. A similar language
change apparently occurred in Western Siberia where the Paleo-
Asiatic inhabitants were linguistically assimilated by the Uralicspeakers,
giving origins to the Samoyeds.
The introduction of food production changed the linguistic
and genetic landscape of the Old World. Colin Renfrew (1987,
1992) has proposed that this subsistence shift was associated
with the spread of Indo-European languages from Anatolia over
most of Europe. Cavalli-Sforza et al. (1994) have demonstrated
with synthetic maps of principal component scores computed
from gene-frequency data that there was a population expansion
(demic diffusion) from Anatolia into Europe, which made
especially Southern- and Central Europeans genetically closer to
the Near Eastern populations. The archeological evidence (for
example, in Zvelebil 1986) indicates that the spread of farming
into northern Europe was slower and took place primarily
through cultural diffusion. As a result, North European gene
pools received less Near Eastern genes than those of Centraland
Southern Europeans and the non-Indo-European languages
survived longer. The early historical period Pictish language of
Scotland and the Finno-Ugric languages of northern and
eastern Europe represent remnants of indigenous North
European languages.
The earliest farming in the East Baltic region dates to the
Corded Ware (Battle/Boat Axe) period (3200-2500 cal. BC)
(Lang 1998). The spread of this culture over this region must
have had a significant impact on the ethnogenesis of the Baltic-
Finns. For instance, the Baltic-Finnic protolanguage is thought
to have received its Baltic loan words as a result of this event.
Central European genes arrived also. Sajantila and PŠŠbo (1995)
have even proposed that the Indo-European-speaking Battle Axe
people became linguistically assimilated by the indigenous
Uralic-speaking people (Lapps/Saami) in Finland. This theory is
based on observations that the Finns are genetically rather
lingistically European, while the Saami are very distinct.
Sajantila and PŠŠbo’s (1995) theory has three problems.
First, it is unlikely for numerically superior intruders with a
more complex social organization to become linguistically
assimilated by numerically inferior indigenous people with a
simpler social organization. Second, the Estonians and the Livs,
living south of the Gulf of Finland, are Finno-Ugrians, although
there have never been Lapps in these regions who could have
linguistically assimilated the Indo-Europeans. Third, the socalled
Lapps who lived in southern and central Finland during
the Middle Ages may have been genetically similar to the
modern Finns and distinct from the modern Saami of northern
Fennoscandia. For these reasons, I consider it more likely that
the Battle-Axe people who arrived in Finland spoke a Proto-
Baltic-Finnish language (Uralic language influenced by Proto-
Baltic languages) and that they were not markedly different
genetically from the inhabitants of southern Finland. This
theory is supported by archeological evidence from Estonia,
indicating the Estonian Corded Ware culture had an indigenous
background (Lang 1998). If this was indeed the case, the Baltic-
Finnic language phylum and people emerged in Latvia and
Estonia around 3200 cal. BC, when local hunter-gatherers came
under considerable linguistic and cultural influence of farming
populations living further south. The appearance of the Corded
Ware culture in Finland would then indicate the arrival of the
Baltic-Finns in Finland around 3200 cal. BC.
A recent estimation of the date of genetic divergence of the
Estonians and Finns from the mitochondrial DNA data might
support this theory. Sajantila et al. (1996) estimated a date of
about 3900 (1900 BC) years for the arrival of the Finns in
Finland assuming the mutation rate of one substitution in 50
generations and a generation time of 20 years. If the average
generation time is 25 years (which is more probable), this date
was 4875 cal. BP (2875 cal. BC), which agrees well with the
arrival of the Battle Axe (Corded Ware) culture in Finland.
However, if the Estonian-Finnish divergence date is based on
average mutation rates across the entire nucleotide sequence
and the mutation rate is one substitution in 5,000 years
(independent of the number of generations), this divergence
occurred 10,000 cal. BP (8000 cal. BC) when Finland was first
settled. I will return to these dates ahead.
Whether the genetic ancestors of the Finns arrived in
Finland 5,000 or 10,000 years ago, the divergence of the Baltic-
Finns into different ethnic groups either started or accelerated
during the Bronze Age (1500-500 BC). The Scandinavian
Bronze culture arrived in the southern and western coastal
regions of Finland with Scandinavian immigrants, who had a
lasting effect on Finnish gene pools and language. The Finns
became genetically similar to the Scandinavians and received
Proto-Germanic loan words. These early Scandinavian
immigrants were linguistically and genetically assimilated into
the indigenous population.
The Middle Ages brought more Scandinavian immigrants
into Finland. The Swedish-speaking minority of southern and
western coastal regions is descended from the Swedish
immigrants who arrived during the years AD 1100-1300. This
immigration reduced the Finns’ genetic distance from the
Swedes (possibly at the expense to that to the Estonians)
because there was from the beginning a considerable amount of
intermarriage (Virtaranta-Knowles et al. 1991).
The linguistic relationship of the Baltic-Finnish and Saami
languages has led to an assumption that the Baltic-Finns and the
Saami descend genetically from the same population, and the
Baltic-Finns’ ancestors diverged from the Saamis’ ancestors as a
result of fairly recent Baltic and Germanic influences on the first
mentioned (see HŠkkinen 1996 for a review). However, the
Baltic-Finns and Saami are genetically too distant from each
other to have descended from the common ancestral
population that lived only about 5,000 years ago. Therefore, it is
more likely that ancestors of the Finns and Saami separated
from each other at the beginning of the Last Glacial Maximum
about 23,000 BP and met each other again in northern
Fennoscandia around 8500 BP.
If the above argument is correct, the so-called Lapps who
lived in southern and central Finland still during the early
historical period and the modern Saami of northern
Fennoscandia were genetically distinct from each other.
Archeological evidence may support this theory. Matti Huurre
(1983) has argued that the southern boundary of the real Saami
was located near the north-south midline of Finland. The
genetic distinctness of the modern Saami of northern
Fennoscandia (for instance, the “Saami-specific motif” in their
mtDNA discovered by Sajantila et al. 1995) may have been
inherited from the earliest inhabitants of the Arctic coast, who
had arrived in the north along the Norwegian coast. These
populations, whose ancestral language is unknown, could have
become Uralic (more specifically Lappic)-speakers as a result of
cultural contacts with Proto-Lappic speaking people arriving
from the south. If this was the case, the Lapps of the southern
half of Finland would have spoken Lappic (Saami) dialects, but
were genetically similar to populations living further south in
the East Baltic region. These southern Lapps were also
incorporated into the gene pool of northward expanding
Finnish populations during the Middle Ages. The reason why
their genetic contribution to the gene pool of the Finns is not
detectable may be their genetic similarity with the assimilators.
The genetic affinities of the so-called Lapps of the southern
half of Finland are key to solving the Finns’ origin. Before we
know the genetic structure of the so-called Lapps of the
southern half of Finland (through extraction of DNA from old
bones) we will never be able to determine the origins of the
Baltic-Finns and when the Finns arrived in Finland. If the
southern Lapps were genetically Saami, the Finns’ ancestors
probably did not arrive in Finland until about 3200 BC. If these
southern Lapps were genetically non-Saami, the Finns’ ancestors
could have arrived in Finland when the region received its first
inhabitants about 10,000 years ago. In light of the available
evidence, we can be certain, however, that the Finn’s ancestors
arrived in Finland at least 3,000 years earlier than the traditional
migration theory would allow.
Conclusion
Whenever the Finns’ ancestors arrived in Finland, we cannot
deny the European origin of the Baltic-Finns and other Finno-
Ugric people based on available physical anthropological and
archeological data. The genetic and possibly linguistic ancestors
of most of the Finno-Ugrian people were very likely the huntergatherers
who inhabited the periglacial zone located between
the Carpathian Mountains and the Volga River during the last
glacial maximum. We could say that from a purely physical
anthropological point-of-view, the Baltic-Finns are either the
easternmost Northwest Europeans or the westernmost Northeast
Europeans.

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