ETRUSCANS, VENETI
and Slovenes:
A
Genetic Perspective
J. Skulj
The Hindu Institute of Learning,
Toronto, Canada. 2005-4-22
INTRODUCTION
In 1973, in
his book Before Civilization: The Radiocarbon Revolution and Prehistoric
Europe, well known archaeologist and pre-historian Lord Colin Renfrew wrote:
"The study of prehistory today is in a state of crisis. Archaeologists
all over the world have realized that much of pre-history, as written in existing
textbooks, is inadequate, some of it quite simply wrong........the chronology
of prehistoric Europe betrays a serious flaw in archaeological theory....Most
of us have been brought up to believe, for instance, that the Pyramids of Egypt
are the oldest-stone built monuments in the world, and that the first temples
built by man were situated in the Near East...... There, it was thought metallurgy
was invented.....It comes, then, as a shock to learn that all this is wrong.
The megalithic chamber tombs of western Europe are now dated earlier than
the Pyramids...... Copper metallurgy appears to have been underway in the Balkans
at an early date-earlier than in Greece...
Already, twenty
years ago, the new scientific technique of radiocarbon dating brought archaeologists
several surprises. But it did not challenge the basic assumptions underlying
what they had written: the position of the ancient civilizations of Egypt and
Mesopotamia as the innovators, illuminating the rest of the Old World with the
radiance of their culture was not challenged. Today the second radiocarbon
revolution, based on recent tree-ring dating, has undermined these assumptions.
Indeed, it is bringing down the whole edifice of links and connections
that were so laboriously built up by scholars over the last fifty years in order
to date and make intelligible our prehistoric past." (Renfrew 1973)
"In reconstructing
the past it is natural to look for clues in stone tools, pottery and grave goods.
These are the tangible artifacts of the past-but they are not the only
survivors. In every cell of our bodies we all carry DNA which has been
passed down almost unchanged from our earliest ancestors...DNA (deoxyribonucleic
acid) is the messenger of heredity" (Sykes & Renfrew 2000).
The mtDNA and
Y chromosome have the potential to be particularly informative for the studies
of human migrations that populated Europe. Both components of the human
genome are inherited from only one parent. Therefore, neither can recombine,
and thus both change by the accumulation of sequential mutations along radiation
lineages. Since all the sequence variants of the mtDNA or Y chromosome
remain associated with each other in total linkage disequilibrium, the sum of
the sequence variant sites of mtDNA or Y chromosome are designated its "haplotype".
Groups of haplotypes sharing distinctive sequence variants inherited from
a common ancestor are known as `haplogroups" (Lell & Wallace 2000).
Analysis of
maternally inherited mitochondrial DNA (mtDNA) of modern populations has become
a useful tool for human population studies and for reconstructing aspects of
evolutionary history. The maternal mode of inheritance of the mtDNA, allows
it to be used for inferring the pattern of prehistoric female migrations and
peopling of different regions of the world. It is now technically possible
to validate these analyses by directly studying the DNA of ancient people and
comparing them to the present day populations (Malyarchuk 2003, Vernesi 2004).
Table 1-The
Age and Frequency of the mtDNA Haplogroups and Subhaplogroups:
|
|
B |
B |
V |
S |
P |
R |
S |
E |
I |
|
|
|
S |
S |
E |
L |
O |
U |
W |
U |
N |
|
|
|
K |
K |
N |
O |
L |
S |
E |
Avg. |
I |
|
|
|
1 |
2 |
|
|
|
|
|
|
|
|
|
|
(I) |
(MM) |
(MP) |
(M) |
(M) |
(M) |
(T1) |
(R) |
(K) |
|
Hg |
Age |
(R) |
% |
% |
% |
% |
% |
% |
% |
% |
% |
H |
19
000-21 |
400
|
37 |
50(T3) |
41 |
47 |
45 |
42 |
41 |
46 |
3 |
HV* |
20
700-22 |
800 |
na |
na |
1.5 |
0 |
1 |
2 |
na |
na |
na |
pre*V
|
10
300-15 |
100 |
na |
0(T2) |
1.5 |
3 |
5 |
5 |
na |
na |
na |
preHV |
15
000-42 |
000 |
na |
0 |
na |
0 |
0 |
0.5 |
na |
0.3 |
na |
V
|
11
000-17 |
000 |
0 |
12(T2) |
4 |
4 |
na |
4 |
5 |
5 |
0 |
J
|
22
000-27 |
000 |
12 |
2 |
10 |
10 |
8 |
8 |
3 |
9 |
0 |
T*
|
33
000-40 |
000 |
8 |
2 |
19 |
5 |
9 |
9 |
3 |
9 |
1 |
T1
|
6
000-13 |
000 |
na |
2 |
3 |
1 |
2 |
2 |
na |
2 |
na |
K
|
13
000-18 |
000 |
20 |
3 |
3 |
4 |
3 |
3 |
14 |
6 |
0 |
U |
(Total) |
|
17 |
(15) |
(7) |
(20) |
(15) |
(16) |
16 |
(16) |
(23) |
U1
|
2
400-52 |
000 |
na |
na |
3 |
0 |
0 |
1 |
na |
0.5 |
na |
U2
|
23
000-48 |
000 |
na |
1 |
0 |
1 |
1 |
1 |
na |
0.6 |
na |
U3
|
11
900-26 |
800 |
na |
0 |
1.5 |
2 |
0.5 |
1 |
na |
1 |
0 |
U4
|
16
100-24 |
700 |
na |
0 |
0 |
6 |
5 |
3 |
na |
3 |
na |
U5
|
45
100-52 |
800 |
na |
12 |
1.5 |
11 |
9 |
10 |
na |
9 |
na |
U6 |
|
na |
na |
1 |
0 |
0 |
0 |
0 |
na |
na |
0 |
U7
|
11
000-45 |
400 |
na |
1 |
0 |
0 |
0.2 |
0 |
na |
0.2 |
na |
I
|
27
200-40 |
500 |
0 |
0 |
3 |
2 |
2 |
2 |
0 |
2 |
2 |
W
|
17
100-28 |
400 |
0 |
1 |
0 |
5 |
4 |
2 |
0 |
2 |
0 |
X
|
17
000-30 |
000 |
1 |
1 |
4 |
1 |
2 |
3 |
0 |
1 |
0 |
Other |
|
|
5 |
na |
2 |
0 |
3 |
2 |
na |
0.3 |
80 |
Hg
Avg. |
(x1000
yrs.) |
|
|
|
|
24.4 |
|
|
|
24.6 |
|
The columns
do not add up to 100%, because the data are from many sources.
Abbreviations:
Hg means haplogroup; (20) represents the total of the subhaplogroups; BSK1 is
the mtDNA obtained from 4,000-5,000 year old skeletal remains from the Basque
region; BSK2 are present day Basques; VEN are Italians from Veneto; SLO are
Slovenes; POL are Poles; RUS are Russians; SWE are Swedes; EU Avg. is European
average; IND are East-Indians.
Using the data
of (R) (Richards 2000), (I) (Izagirre 1999), (MM) (Maca-Mayer 2003), (MP) (Mogentale-Profizi
2001) and summarized in Table 1, a weighted average haplogroup age has been
calculated. The weighted average haplogroup age of Europeans as a whole
is~ 24,600 +/-2,800 years, but the Slovenian haplogroup age is almost the same
age at ~24,400 +/- 2, 700 years. older at 27,460 years. The reason for
the old haplogroup age in Slovenes is a relatively high frequency of haplogroup
U, which is 25 % more frequent in Slovenes than Europeans as a whole.
The Origins
and Diffusion of Haplogroups:
Haplogroup
H is the most common haplogroup in Europeans. It is also common in Caucasoid
populations of the Near East and North Africa and is also observed in northern
India. Even though this haplogroup is more common in Europe than in the
Near East, its diversity is much higher in the Near East than in Europe and
this suggests that, haplogroup H originated in the Near East ~25,000 years ago
and then expanded into Europe ~20,000 years ago (Torroni 1998).
Haplogroup
V is is more recent and is considered to have European or North African origin
~13, 000 years ago (Torroni 1998).
Haplogroup
J has its origin in the Near East and reaches its highest frequency in Arabia
amongst Beduins and Yemeni at 25 % and that it may have accompanied the spread
of farming to Europe (Richards 2000). It is haplogroup J that is associated
with the arrival of farming at the beginning of the Neolithic period (Renfrew
2002).
Haplogroup
T is a sister of haplogroup J with its origin in the Near East ~50,000 years
ago and more recent date in Europe (Richards 2000).
Haplogroup
K is widely distributed amongst the populations of Europe, Near East and India
(Torroni 2000, Kivisild 1999). The 5200 year old `Ice Man' from the Alps,
also carried the haplogroup K maternal lineage (Di Benedetto 2000).
Haplogroup
U is considered the oldest. This haplogroup is present in Slovenian population
at ~ 20 %; this considerably higher than the European average of 16 %. Subhaplogroup
U5, which is part of Hg U is estimated to be the oldest. It is present
in modern Basques at 12 %, Slovenes at 11 % and Indians at <1 %. U5
is to have evolved mainly in Europe ~50 000 years ago (Richards 2000), along
with Hg V at a more recent date (Achilli 2004). This may account for some of
the linguistic similarities between Basque and Slavic languages as noted by
Jandacek and Arko (Jandacek 2001). It should be noted that Tambets et
al. consider U4, which is another subhaplogroup of U, to be even more European
than U5 and note that it is relatively frequent in the Balkans and Eastern Europe.
In Slovenes the frequency of U4 at 6% is double the European average.
Both U5 and U4 are recognized as Upper Paleolithic in age and their beginnings
are before farming; a few U4 lineages have also been found in India (Tambets
2002).
Haplogroup
X is shared between Europeans and American Indians (Torroni 1998). The
estimated arrival from Europe to the area of the Great Lakes is ~15,000 years
ago.
The research
by Richards et al. (1996), based upon the analysis of mitochondrial DNA, suggests
that the greater part of the variability of the mtDNA in Europe is to be dated
back to the Late Upper Paleolithic (70 %), with the significant proportion to
Upper Paleolithic (10 %) and only limited proportion (20 %) to haplogroup J,
associated by them with the arrival of farming at the beginning of the Neolithic
period (Renfrew 2002). Into the area of the Alps, only a relatively small
number (7 %) of lineages have entered since the Bronze Age up to the present
times (Richards 2000).
ETRUSCAN
and VENETIC mtDNA LINEAGES:
Vernesi et
al. obtained fragments of well preserved skeletons from Etruscan necropolises,
covering much of the Etruria in terms of both chronology (7th
to 2nd centuries B.C.) and geography. The tombs typically
belong to the social elites, so the individuals studied may represent only a
specific social group, the upper classes. The ancient human remains came
from the following sites: Adria, Volterra, Castelfranco di Sotto, Castellucio
di Pienza, Magliano and Marsiliana, Tarquinia and also Capua. Two cities,
Adria in the Po valley and Capua in Campania, were at the fringes of Etruscan
territory. In Adria the hybridization with the Veneti may have occurred
(Vernesi 2004).
Vernesi et
al. compared the mtDNA results obtained from the ancient remains to a number
of modern populations. Unfortunately, they did not take into account the
genetic studies of Slovenes ( Malyarchuk 2003), who are geographically relatively
close to Adria.
The Etruscans
are one of the mysterious peoples of the ancient world, who seem to have appeared
for a time on the stage of history, and then seemed to have disappeared. In
fact, from the end of the Roman period to the Middle Ages, they could be said
to have ceased to exist, since the sites of their cities, towns, villages and
farms had been completely lost. It was in the19th century that the study
of the Etruscan legacy began in earnest. The heart of Etruria was the
territory, in the present day Italy, on the Tyrrhenian Sea between the
rivers, Arno on the north and Tiber on the south and extending to Perugia in
the east. The Etruscan influence in the 7th and
6th centuries B.C., went beyond its heartland and extended
to, Adria in the Po valley in the north and to Capua in the south. It
is generally accepted, that present day Tuscans are the Etruscans' closest neighbors
(Wellard 1973, Vernesi 2004).
The Veneti
are also one the historic peoples, subject of many discussions and debates,
but who were more widespread than the Etruscans. They were present in
many lands (Mogentale-Profizi 2001): Veneti in Paphlagonia -northern coast of
present day Turkey-were mentioned by Homer in 9th cent.
BC., Veneti in Illyricum (Enetoi) on the lower Danube and in the upper
Adriatic, were mentioned by Herodotus in 5th cent. BC:,
Veneti in central Europe mentioned by Tacitus and Pliny the Elder, Veneti in
Gaul were mentioned by Caesar, and Veneti in Latium who are referred to as Venetulani
by Pliny the Elder. The Veneti and Etruscans appear to be related. However,
Adria was in the 10th Roman province `Venetia et Histria'
until the downfall of the empire. There is historical, linguistic and
topographic evidence that present day Slovenes are indigenous to their land
and descendents of the Veneti (Šavli 1996).
DISCUSSION
of GENETIC STUDIES
In the bone
fragments, taken from the tombs of Etruria, Capua and Venetia, Vernesi et al.
(2004) have found that haplogroup H was the most common; other haplogroups such
as JT and preHV were also present. When they made a more refined comparison
using the haplotypes that are components of haplogroups, they found that out
of 22 mtDNA HVS1 haplotypes, which they observed in 28 skeletal remains, only
two of them, CRS and 16126, occur in a sample of modern Tuscans and are carried
by ~14% of them. Tuscans are considered to be the descendants of the Etruscans.
Both haplotypes occur in skeletons from Adria and Magliano/Marsiliana.
The fragments from Magliano/Marsiliana have been dated at 7th-6th
centuries B.C., whereas those from Adria are from 5th-4th
centuries B.C. (Vernesi 2004).
Table 2-
Mt DNA Comparison of Etruscans, Veneti and Slovenes for HVS 1 Haplotypes:
Place |
Age |
Hg |
HVS1 |
Slo.HVS
1 |
Hg |
Frequency (%) |
Adria |
2450 |
H |
CRS |
CRS |
H |
12 |
(5) |
|
H |
16223 |
16223 |
H |
1 |
|
|
H |
16129 |
16129,16304 |
H |
1 |
|
|
|
16126 |
16069,16126 |
J* |
8 |
Volterra |
2450 |
H |
16261 |
16261 |
H |
1 |
(4) |
|
|
|
|
|
|
Mag/Mar |
2650 |
H |
16311 |
16311 |
H |
2 |
(6)
|
|
H |
CRS |
CRS |
H |
12 |
Comparing the
results of Vernesi et al and Malyarchuk et al, it becomes apparent that, the
present day Slovenes, carry more than just CRS and 16126 `Etruscan' mtDNA
HVS1 haplotypes found in the Tuscans. Twice as many `Etruscan' haplotypes
have been found in Slovenes than in Tuscans, namely: CRS, 16261, 16223, 16311.
These were found in skeletal remains from Adria, Magliano/Marsiliana and
also from Volterra. Two additional haplotypes from Adria, 16126
and 16129, are similar to Slovenian haplotypes, but the Slovenian haplotypes
differ from the `Etruscan' of Adria, by one substitution each, namely 16126
with 16069 and 16129 with 16304. However, haplotype 16129 without the
16069 substitution is found in Bosnia. This leaves just one haplotype
out of five, namely, 16126-16193-16278, where no similar haplotype is found
in Slovenia. However, this 16126-16193-16278 haplotype is similar
to that found in skeletal remains from Capua at the southern limit of Etruscan
influence where hybridization with Samnium natives or Greek colonizers may have
occurred (Malyarchuk 2003, Vernesi 2004).
The root type
16069-16126 HVS1 sequence, present in ~8% of Slovenes, is very diverse and
may represent a trace of Neolithic (new Stone Age at the beginning of agriculture)
migration from the Middle East (Malyarchuk 2003). Haplotypes CRS,
16223, 16261 and 16311 are carried by ~17% of Slovenes. They belong
to haplogroup H, which is estimated to be ~20, 000 years old; this haplogroup
is the most common one in Slovenes at 47% (Richards 2000, Malyarchuk 2003).
Ancient
Human Remains from Adria in Veneto
Focusing on
5 haplotypes, CRS, 16126, 16129, 16223, 16126-16193-16278 found in skeletal
remains from Adria, which was part of Venetia et Histria during the Roman era,.(Adria
is even now located in Veneto, Italy), and comparing them to the present day
populations, we find:
--CRS in Slovenes
at 13% (Malyarchuk 2003), in Europe at 24% (Richards 1996)
--16126 is
found as 16069-16126 in Slo at 8% (M), in Eu 16069-16126 is at 7% (R)
--16129 is
found in Bosnians (Bos) at 2% (M), in Russians at 1% (M1) in Basques at
9% (R); in Slo it is found as 16129-304 at 1% (M).
--16223 is
found in Slo at 1%, elsewhere in Eu only in South Germans and Ukrainians at
11 % (M).
It is noteworthy
that 4 out of 5 or 80 % of the `Venetic' sequences are found in Slovenia, where
these sequences represent 23 % of the extant population. It is also significant
that Richards et al., (2000), in their study of 520 individuals from across
Europe, (for the Slavic populations only Bulgarians, Czechs, Poles and Russians
were included in their study), did not detect the 16223 haplotype, which is
present in skeletal remains from Adria, nor has it been found in a sample of
modern Tuscans (Richards 1996, Vernesi 2004), but is has been found only in
Slovenia, South Germany and Ukraine (Malyarchuk 2003).
In addition
to the haplotypes found in the ancient Veneti from Adria, Slovenes also share
haplotypes with the skeletal remains of Etruscans from Etruria proper, namely
from Volterra (Vo) and Magliano/Marsiliana (M/M). Furthermore, Russians
and Poles share one lineage with Castelfranco di Sotto (CS) not found in the
Slovenian sample.
--16261 of
Vo is found in Slo at 1% (M), in Eu at <1% (R).
--16311 of
M/M is found in Slo at 2%, in Bosnians at 7% (M), in Eu at 5% (R)
--16126 of
M/M is found in Slo as 16069-16126 lineage at 8% (M) in Eu at 7% (R).
--CRS of M/M
is found in Slo at 13% (M), in Eu at 24% (R).
--16189-16356
of (M/M) is found in Poles at 0.5%, Russians at 0.5% and Germans at 0.4% (M1)
Here again,
there is no abrupt genetic differences between skeletal remains from Etruria
proper and the present day Slavic populations in the Balkans, since 27 % of
Slovenes share Etruscan sequences.
From the above
comparison, it can be seen, that there is a genetic continuity between ancient
populations as attested from the skeletal remains found in Etruria proper and
especially between those found in Venetia and the present day Europeans. While
Tuscans share 2 haplotypes with the Etruscans, Slovenes and Bosnians share
3 haplotypes. It should also be noted that 2 additional Etruscan haplotypes
from Adria in Veneto, differ from the Slovenian haplotypes by one substitution.
Considering the evidence, this shows the relatively strong genetic mtDNA
relationship between ancient Veneti and modern day Slovenes.
Alpine Prehistoric
Human Remains:
Previously
Di Benedetto G, et al. (2000), after rigorous controls to ensure their authenticity,
published mtDNA sequences from 3 prehistoric human remains from the Alps between
6000 and 14000 years of age.
Table 3-Comparison
of mtDNA of 3 Prehistoric Humans from Alps and Slovenes:
Place |
Age |
Hg |
Haplotype |
Slovenian Haplotype |
Hg |
Mezzocorona |
6400 |
T |
16126,16292, 16294 |
16126,
16294, 16296, 16304 |
T |
Villabruna |
14000 |
H |
16261,
16274 |
16261 |
H |
|
|
|
|
16274 |
H |
Borgo
Nuovo |
6000 |
H |
CRS |
CRS |
H |
|
|
|
|
|
|
The authors
conclude after also taking into consideration the sequence of the 5200 year
old `Ice man' belonging to haplogroup K,(which is also present in Slovenes
at ~4 %) that these sequences provide some insight into prehistoric European
mtDNA diversity and that mtDNA diversity in Europe was not much different
in the Neolithic period than it is at present. Each Neolithic sequence
falls into a different haplogroup, testifying to the high level mtDNA diversity
in the Alps at the beginning of the Neolithic period. The fact that haplogroups
that are common in Europe today are also found at the beginning of the Neolithic
period further supports a genealogical continuity in Europe between the Neolithic
and the present (Di Benedetto 2000).
Y chromosome
Perspective:
What language
did the Etruscans and/or Veneti speak? Barbujani has made an intriguing
observation, that partial correlations with language are stronger for the Y
chromosome than for mtDNA (Barbujani 1997). Conventional opinion has it,
that Etruscans spoke a language isolate, a non-Indo-European language and that
it disappeared ~90 B.C., when they lost their autonomy to the Romans (Vernesi
2004).
Some Slovenian
scholars held/hold a different view. Bor had postulated that Etruscans
were people originally linguistically related to the Veneti; (the genetic evidence
supports his hypothesis); they came from the north and in course of time merged
with another people, which in turn influenced their language. By using
Slavic languages, as a point of reference, he was able to decipher some of the
older Etruscan inscriptions, including the Pyrgian Tablets, but not their later
inscriptions. On the other hand, he was quite successful in deciphering
the Venetic inscriptions (Šavli 1996).
The Y chromosome
studies reveal that Haplogroup I (Hg I), reached ~40%-50% in two distinct regions-in
Nordic populations in Scandinavia and around the Dinaric Alps. Overall,
this suggest, that populations carrying the Hg I could have played a central
role in the process of human re-colonization of Europe, after the Ice Age (Rootsi
2004). Semino proposes that Hg I (M170) haplogroup originated in
Europe in descendants of men that arrived from Middle East 20,000 to 25,000
years ago. This can be associated with an Epi-Gravettian culture in the
area of the present-day Austria, the Czech Republic and the northern Balkans
(Semino 2000).
Subhaplogroup
HgI1b* is the most frequent clade in eastern Europe and the Balkans; its subclade
Hg I1b2 is found in Sardinia, Castille and in Basques (6%). Rootsi et
al., make an observation and also show graphically, that Hg I1b* and Hg I1b2
co-occur west of the Italian Apennines. However, in the Veneto region
of Italy Hg I b2 is absent, but Hg I1b* occurs at a frequency of~10% and only
Hg I1b* is present west of the Apennines. The frequency increases towards
the east. East of the Adriatic Hg I1b* reaches its highest concentration
in the north western Balkans (Rootsi 2004). This is also another indication
that, there is also a genetic continuity, between the Slovenes and the people
of Veneto region, including Adria. This genealogical continuity is based
on the paternally inherited Y chromosomes. This is in addition to the
maternally inherited mtDNA, as discussed earlier.
Barbujani in
his paper ``Genetics and the population history of Europe'', shows graphically
a genetic continuity between the populations of the north western Balkans and
the peoples now occupying the land of the ancient Veneti and Etruscans in Italy.
A clear demarcation is seen in northern Italy at the western boundary
of the Veneto region (Barbujani 2001). In another genetic study of the
present day populations, it has been found, that the population in eastern Veneto,
is more akin to Tuscanian, than to western Veneto population (Mogentale-Profizi
2001). Furthermore, Malyarchuk et al., have also noted, that Slovenes
have a high frequency, at 5%, of H-subcluster 16162, which is characteristic
for central and eastern European populations. In the western neighbors
of Slovenes, in the present day Veneto speakers of Italy, this is also present,
at a similar frequency of 6% (Malyarchuk 2003).
CONCLUSION
Genetic information
indicates that inhabitants of Slovenia are genetically very ancient population.
There is evidence
of genealogical continuity between 5,200-14,000 year old skeletons from the
Alps, the ancient Etruscans and Veneti and the present day Slovenes.
Genetic information
makes it evident, that Slovenes are indigenous to their land as indicated
by the mtDNA relationship with the ~2,500 year old skeletal remains of the Etruscans
and Veneti, particularly those from Adria. The Y-chromosome DNA data on
the extant populations also support this premise.
Genetic evidence
supports the historic quotation from the biography of St. Columban written in
615 A.D. and cited by Tomažic "Termini Venetiorum qui et Sclavi dicuntur"-the
land of the Veneti who are also called Slavs (Šavli 1996).
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