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ETRUSCANS, VENETI and Slovenes: A Genetic Perspective
J. Skulj
The Hindu Institute of Learning,
Toronto, Canada. 2005-4-22



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).

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).

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
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).

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).


     Barbujani G (1997) DNA Variation and Language Affinities.  Am. J. Hum. Genet. 61:1011-1014.
     Barbujani G, Bertolle G (2001) Genetics and the population history of Europe.  PNAS vol. 98     no.1:22-25.
     Di Benedetto G, et al. (2000)  Mitochondrial DNA sequences in prehistoric human remains from     the Alps.  European Journal of Human Genetics 8: 669-677.
     Jandacek P, Arko L (2001)   Linguistic Connections between Basques and Slavs (Veneti) in Antiquity.  ZBORNIK prve mednarodne konference: Veneti v etnogenezi srednjeevropskega prebivalstva/PROCEEDINGS of the First International Topical Conference: Veneti within the Ethnogenesis of the Central-European Population eds. Perdih A, Rant J Ljubljana, Slovenia: Jutro pp.151-166   ISBN 961-6433-06-7
     (K) Kivisild T, et al. (1999)   Deep common ancestry of Indian and western-Eurasian mitochondrial DNA lineages.  Current Biology 9:1331-1334.
     Lell TJ and Wallace DC (2000)  The peopling of Europe from the Maternal and Paternal and Perspectives. Am. J. Hum. Genet. 67: 1376-1381.
     (M) Malyarchuk BA, Grzybowski T, Derenko MV, Czarny J, Drobnic K, Miscicka-Sliwka D (2003)     Mitochondrial DNA Variability in Bosnians and Slovenes.  Ann. Hum. Genet. 67: 412-425.
     (M1) Malyarchuk BA, Grzybowski T, Derenko MV, Czarny J, Wozniak M, Misicka-Sliwka D     (2002)  Mitochondrial DNA in Poles and Russians.  Ann. Hum. Genet. 66:261-283.
     (MP)Mogentale-Profizi N, Chollet L, Stevanovitch A, Dubut V, Poggi C, Pradie MP, Spadoni     JL,Gilles A, Beraud-Colomb E (2001)  Mitachondrial DNA sequence diversity in two     groups of Italian Veneto speakers from Veneto.  Ann. Hum. Genet. 65: 153-166.
     Renfrew C (1973) Before Civilization: The Radiocarbon Revolution and Prehistoric Europe.

    Great Britain, 1999 Pimlico edition: pp.15-17   ISBN 0-7126-6593-5..

     Renfrew C (2002) `The Emerging Synthesis': the Archaeogenetics of Farming/Language Dispersals and other Spread Zones. in Examining the farming/language dispersal hypothesis. eds. P .Bellwood & C. Renfrew, Cambridge: McDonald Institute for Archaeological Research, 3-16  ISBN 1-902937-20-1  
     Richards M, Macaulay V, Hickey E and 34 others (2000)  Tracing European Founder Lineages     in the Near Eastern mtDNA Pool.  Am. J. Hum. Genet. 67:1251-1267.
     Richards M, Corte-Real H, Forster P and 7 others (1996)  Paleolithic and Neolithic Lineages in     the European Mitochondrial Gene Pool. Am. J. Hum. Genet. 59:185-203.
     Rootsi S, Magri C, Kivisild T and 42 others (2004  Phylogeography of Y-Chromosome Haplogroup I Reveals Distinct Domains of Prehistoric Gene Flow in Europe.  Am. J. Hum. Genet. 75:128-137.
     Šavli J, Bor M, Tomažic I, trans. Škerbinc A (1996) VENETI: First Builders of European     Community.  Wien,Boswell: Editiones Veneti  pp.80, 197-199,344, 443, 501  ISBN 0-    9681236-0-0
     Semino O, Passarino G, Oefner PJ and 14 others (2000) The Genetic Legacy of Paleolithic     Homo sapiens sapiens in Extant Europeans: A Y Chromosome Perspective.  Science     vol. 290 10, November.
     Sykes B, Renfrew C (2000) Concepts in Molecular Genetics. Archaeogenetics: DNA and the     population prehistory of Europe, in Renfrew & Boyle, Cambridge, McDonald     Institute for Archaeological Research, pp. 13-21  ISBN: 1-902937-08-2.
     Tambets K, et al. (2002)  Complex Signals for Population Expansions in Europe and Beyond. Examining the farming/language dispersal hypothesis, in Bellwood & Renfrew, Cambridge, McDonald Institute for Archaeological Research, pp. 449-475  ISBN 1-902937-20-1  
     (T1) Torroni A, et al. (1996) Classification of European mtDNAs from an Analysis of three

    European populations.  Genetics 144:1835-1850.

     (T2) Torroni A, et al. (2001)  A Signal, from Human mtDNA, of Postglacial Recolonization in     Europe. Am. J. Hum. Genet. 69:844-852.
     (T3) Torroni A, et al. (1998)  mtDNA Analysis Reveals a Major Late Paleolithic Population     Expansion from Southwestern to Northeastern Europe. Am. J. Hum. Genet. 62:1137-1152    
     Vernesi C, Caramelli D, Dupanloup I, Bertorelle G, Lari M, Capellini D, Moggi-Cecci J,     Chiarelli B, Castri L, Casoli A, Mallegni F, Lalueza-Fox C, Barbujani G (2004)  The     Etruscans: A Population-Genetic Study.  Am J Hum Genet 74: 694-704.
     Wellard  J (!973)  The Search for the Etruscans. London: Sphere Books Ltd. pp.11-13   ISBN 0     351 18677 8