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Contents: The origin of haplogroup I1-M253 in Eastern Europe, by ALEXANDER SHTRUNOV. Arabian clusters of haplogroup E1b1b1c1 (M34), by AKPER ALIEV, DMITRY TARTAKOVSKY. Origin, Distribution and Migrations of I2b*-Subclades, by HANS DE BEULE. Origins of Hg I-L38 (I2b2) Subclades, by HANS DE BEULE. Early Bronze Age Origin and Late Iron Age (La Tene) Migrations of I-L38, by HANS DE BEULE. Phylogenetic Relations and Geographic Distribution of I-L38 (aka I2b2, by HANS DE BEULE. Haplogroups E1b1b1c1 (M34) and E1b1b1c1a (M84) among Jews. Could Abraham be E1b1b1c1 or E1b1b1c1a? By AKPER ALIEV, DMITRY TARTAKOVSKY. About the influence of population size on the accuracy of TMRCA estimation, done by standard methods using STR locus complex, by DMITRY ADAMOV, SERGEY KARZHAVIN (Translate by VASILI GAVRILOV).

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Page 1: Russian Journal of Genetic Genealogy. Vol 1, №2, 2010

Vol 1, №2, 2010

Page 2: Russian Journal of Genetic Genealogy. Vol 1, №2, 2010

ISSN: 1920-2989

Russian Journal of Genetic Genealogy

Publisher Lulu inc., 2010

All rights reserved. No part of this publication may be reproduced, altered in

any form or by any means: mechanical, electronic, with photocopying, etc., with-

out the prior written permission of the publisher of the journal, or authors of ar-

ticles.

When citing a reference to this publication is required.

Editor

Michael Temosh

Art editor

Nataliya Zyryanova

Technical editor

Denis Grigoriev

Reviewer

Alexander Kireev

Zhaxylyk Sabitov

Dmitriy Adamov

Contact address

[email protected]

© RJGG, 2010

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The Russian Journal of Genetic Genealogy: Vol 1, №2, 2010 ISSN: 1920-2989 http://ru.rjgg.org © All rights reserved RJGG

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The origin of haplogroup I1-M253 in Eastern Europe

Alexander Shtrunov

Abstract

One of the actual issues of modern genetic genealogy is the origin of a haplogroup. It is not easy to connect da-

ta on genetics, archeology, linguistics, anthropology and other related sciences. In this paper the author tries to find the root of haplogroup I1-M253 in Eastern Europe.

Haplogroup I1* together with it’s relative sub-

clades I2a* and I2b* is the native European hap-logroup because its frequencies outside Europe are extremely small. This spreading of carriers of this haplogroup gave basis to consider carriers of I1 as the descendants of Paleo-European popula-tion. The area of haplogroup I1-M253 is concen-trated mainly in the north of Europe, in the Scan-dinavian countries. There are also local of I1 in England (15,4%) [1], Sicily (up to 18,75%) [2] and in the centre of European Russia (up to 17%) [3]. The presence of carriers of haplogroup I1-M253 at British Isles is associated with the ex-pansion of the Vikings and the Normans, which was proved by historical and genealogical stu-dies, though the ancient migrations are also possible. In Sicily, haplogroup I1-M253 strongly correlates with Norman invasions from the terri-tory of modern France (Normandy, I1 – 11,9%) and the foundation of the Kingdom of Sicily (Sici-lian Kingdom) in 1130. However, the presence of haplogroup I1-M253 with high frequencies in the center of the European part of Russia brings up a lot of questions. E.V. Balanovskaya and O.P. Ba-lanovsky state the following about it in their book «The Russian gene pool of the Russian Plain» [4]:

«Spreading of «Nordic» haplogroup I1a in Russian area is considered quite unexpected (Fig. 1). The high values of I1a would be predicted close to Scandinavia in the northwest of Russian area. There, as well as at the western boundary «Varangian» influence can be expected in the form of high frequencies of I1a. However, com-pact maximum of I1a (11-12%) is located in an entirely different area at the north-east. This lo-cal centre stands out against the background of low frequencies (less than 6%), which are typical of the rest of the Russian area. The presence of this center is based on data of three Russian populations studied by extensive sampling. Of course, in comparison with frequencies of I1a in Scandinavia (25-40%), this local maximum is minor. But its remoteness from the main zone of high frequencies of this haplogroup in Scandina-via requires explanation. It is hard to explain this local maximum by close relations between Scan-dinavia and regions of Transvolga and basin of Vychegda excluding other relative Russian re-gions. Apparently, history of population of haplo-group I1a is more complicated than a simple ex-pansion from Scandinavia, and it may include an-cient relations between the Finno-Ugric peoples of Eastern Europe and the ancestors of German-speaking Scandinavians».

____________________________________________________________ Received: May 11 2010; accepted: May 12 2010; published: May 16 2010 Correspondence: [email protected]

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Fig. 1. Map of presence of haplogroup I1 (Balanovsky et al. 2008 [5]).

Krasnoborsk, Archangelsk region 12,1% Vologda 11,6% Unzha, Kostroma region 11,5%

Complete the data about the region in question from works of other researchers to compare the

spatial distribution of haplogroup I1:

Vologda Region 17,0% (Roewer et al. 2008 [3]*) Archangelsk 14,2% (Mirabal et al. 2009 [6]) Ryazan Region 14,0% (Roewer et al. 2008) Tatarstan 13,0% (Genofond.ru**) Moksha people from Staro-Shayga district of Mordovia 12,0% (Rootsi et al. 2004 [7]) Penza region 12,0% (Roewer et al. 2008) Kostroma 11,3% (Underhill et al. 2007 [1]) Tambov region 10,0% (Roewer et al. 2008) Ivanovo region 10,0% (Roewer et al. 2008)

Data from additional sources confirm the exis-

tence of a local maximum of I1-M253 in the cen-tral part of European Russia. The boundaries of

this local maximum can be seen clearly on the map (Figure 2) developed by the author.

_____________________________________________________________ *Calculation of frequency of haplogroup I1 was carried out by predictor, ** data was taken from the atlas at Genofond.ru)

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Fig. 1. Map of distribution of frequencies of haplogroup I1-M253.

Due to the fact that detailed studies (for) cla-rifying the reason of such high local frequencies of haplogroup I1 on such a wide area still don`t exist, let us try to fill up the gap by analyzing da-ta from published papers, linking data of genet-ics, archeology, linguistics, anthropology and other related sciences.

Since there is no reliable data about historical

migrations from Scandinavia, which could have left such a significant mark on the territory of modern Russia, we will try to consider all availa-ble variants.

Goths of Ermanaric. Ermanaric (died in 376)

was the king of the Goths from the Amali clan. Gothic historian Jordanes wrote about Ermanaric [8]: «Soon after Geberich, king of the Goths, had passed away from human deeds, Hermanaric, the noblest of the Amali, succeeded to the throne. He subdued many warlike peoples of the north and made them obey his laws. Many ancient authors had justly compared him to Alexander the Great. Among the tribes he conquered were the Gol-thescytha, Thiudos, Inaunxis, Vasinabroncae, Me-rens, Mordens, Imniscaris, Rogas, Tadzans, Athaul, Navego, Bubegenae and Coldae». (Here the researchers assume the prototype of enume-ration from Russian Chronicles: Rus, Chud and all languages: Ves, Merya, Mordvins (?)...)

Ermanaric and his Goths probably could not

have made a significant mark on the territory of modern Russia - especially in this region, be-cause the existence of such vast empire is quite doubtful (Fig. 3). Even if this Empire had existed, its age wasn`t long because of the invasion of Huns. Goths are associated with Chernyakhov archaeological culture (III century AD), and therefore we should look for descendants of Goths among the mountain population of the Crimean Tartars (Tata) and the Greeks of Azov region, comparing their haplotypes with the hap-lotypes of population of northern Spain and Got-land.

Varangians. Varangian invasions (IX-XII cc.)

could also made their mark in the gene pool of Eastern Europe, though basic routs of Varangians had passed away from examined areas. The finds related to the Scandinavians point rather at trade relations, than expansion. Therefore the Varan-gians had to be excluded from the list of possible contenders who could have left a significant mark in the gene pool of eastern Europe. Though it`s quite possible that a part of the Normans could have settled and assimilated with the local popu-lation of this region [10].

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Fig. 3. «Empire of Ermanaric» in IV century AD and the assumed itinerary of III-IV centuries AD (by BA Rybakov). a – peoples mentioned in the list of Jordanes; b – the order of the peoples; c – the main areas of Cherniakhov culture in II-IV centuries AD.; d – direction of sea expansions in III century AD; e – direction of Slavic colonization in III-IV centuries AD. [9].

Ancient migrations. Probably, we deal with

ancient migrations on the territory of Eastern Eu-rope, assumed by The Balanovskys. That seems reasonable, given the exclusively European area of haplogroup I-M170.

If haplogroup I1-M253 is linked with the an-

cient population of Europe, it is quite possible that they were the speakers of Paleo-European language.

Researches of Serebrennikov BA [11] have

showed that the ancient substrate toponyms of unknown origin (i.e. non-Uralic and non-Indo-European) are presented on the territory between Volga and Oka rivers. Also it is widely spread in the Nizhny Novgorod area (no data), Chuvashia (7,5% I1), Kirov region (no data), Vologda region (17% I1), Archangelsk region (14,2%), in Karelia (8,6% I1) and in the west of Smolensk region (2% I1).

Examples of substrate toponyms with end-ings:

– ga (Yuzga – branch of Moksha, Arga –

branch of Alatyr, Vyazhga – branch of Moksha and Volga)

– ta (Pushta – branch of Satis, etc.); – sha (Ksha – branch of Sura, Shoksha); – ma (Losma – branch of Moksha, Shalma –

branch of Sivin); – da (Amorda) [12]. On the basis of Serebrennikova’s study Tre-

tyakov PN offered a hypothesis that this Paleo-European language belonged to the creators of the Neolithic cultures of the comb ceramics (CCC) [13].

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CCC had originally occupied the territory of the Volga-Oka-Klyazma rivers. In 3rd millennium BC its carriers moved to the north and north-west, where they settled on the territory from the Baltic sea to Vychegda and Pechora.

Craniological data suggest that the carriers of CCC of Lyalovo type are very heterogeneous. Lyalovo people was generated by alien Nordic population of Sami subrace (Fig. 4, at right) and the aboriginal Mesolithic population of Caucasoid people of Volga-Oka post-Swiderian culture - as the carriers of Upper Volga culture [14].

Fig. 4. Sculptural reconstruction of the skull of a man from Valadar* (the lower reaches of the Oka region) [15]

and the skull of a young man from the burial № 19 of the Sakhtysh II sepulcher (Ivanovo region) [16]. Culture of Upper Volga had spread in the cen-

ter of the Russian Plain since the turn of the 6-5th millennium BC until the end of 5th millennium BC. The earlier monuments of Upper Volga culture

are concentrated in the eastern part of its area, the later – in the central and western areas that is apparently due to the arrival of new alien population.

Fig. 5. Area of Upper Volga, Volosovo and Fatyanovo cultures [17].

_____________________________________________________________ *This site refers to the culture more ancient than Upper Volga and Volosovo archaeological cultures, but the sculptural reconstruction reflects more close-ly the image of Caucasoid population of the Volga-Oka Mesolithic post-Swiderian tradition

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At present the hypothesis of origin of Volga-Oka culture and CCC from the Volga-Oka Meso-lithic post-Swiderian tradition is the most plausi-ble, since the transition from Mesolithic to Neo-lithic was smooth and Butovo culture prevailing in the end of the Mesolithic in the Volga-Oka region also succeeded to the Swiderski tradition [18].

Swiderian culture is the archaeological culture

of the final Paleolithic on the territory of Central and Eastern Europe. Due to the changes of cli-matic conditions in the 11-10th millennium BC people of Sviderian culture began to move from the area of modern Poland, Belarus and Lithuania to the east and reached the given region of Vol-ga-Oka rivers in 8th millennium BC.

The same processes forced the migrations of the neighboring population of carriers of Ahrens-burgian tradition, probably allied to the people of Sviderian culture. This Paleolithic culture existed in 10-9th millennium BC in Denmark and Northern Germany; the main occupation of this population was hunting for reindeer.

In 10th millennium BC people of Ahrensbur-

gian culture began to move following two direc-tions of the retreating ice cover – to the north-west and north-east, passing along both sides of the Baltic glacial lake (Fig. 6).

Fig. 6. Stages of the formation of Baltic Sea basin [19]. 1. Baltic Ice Lake (about 16 thousand years BC). 2. Yoldia Sea (about 7,9 thousand years BC). 3. Ancylus Lake (about 6,8 thousand years BC). 4. Littorina Sea (about 5 thousand years BC).

Arensburgian people established a number of

so-called «culture of Maglemose» in 8-6th millen-nium BC. These are such cultures as Fosna-Hensbacka in Sweden and Norway, Komsa in the far north of Scandinavia, including the Kola Pe-ninsula, Askola and Suomusjärvi in Finland and Karelia, Veretye in east the lakeside of Ladoga, Kunda in the Neva region, Estonia, Latvia and Maglemose in England, Northern Germany and Denmark.

It is necessary to mantion that most high di-versity of haplotypes of I1-M253 - a is fixed it in Denmark [1], which is the starting point of people of Arensburgian culture.

If the assumption about the relationship be-

tween carriers of Ahrensburgian and Swiderian cultures is correct, then we will observe similar anthropological, linguistic and genetic situation in Scandinavia.

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Fig. 7. Mesolithic (8.-5. mil BC) roots of Early Iron Age substratum components. [18].

Legend: 1 – Maglemose-Ertebølle tradition (M – Maglemose incl. English Maglemose; F – Fosna; K – Komsa; A – Askola; S – Su-omusjärvi); 2 – Świdry tradition (Ś – Świdry, Co – Baltic Typical Comb Pottery culture); 3 – area of formation of Pit-and-Comb

Pottery cultures of Central Russia. Anthropological type of carriers of Ahrensbur-

gian culture had a characteristic sharp dolicho-cranic (Fig.8.) broad-faced Caucasoid type, which corresponds well to the post-Sviderian aboriginal type in the Volga-Oka region, who had a dis-tinctly Caucasoid dolichocranic type with the nar-row face. The same anthropological type partici-pated in formation of the Sami. Comparison of anthropological data shows that Sami type

presents features of the ancient North-European population and more recent features associated with penetration of the Mongoloid features to the north. The combination of the two components led to the formation of anthropological features of Sami people. Dentistry also notes the existence of ancient Northern and recent Mongoloid fea-tures among Sami [20].

Fig. 8. Examples of different skull types [21].

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In the Sami language two components are al-so made out. The first is Pre-Finnish substratum, the second is the Old Finnish, which is close to the Baltic-Finnic and Finno-Volgaic languages. The legacy of the Pre-Finnish substratum can be seen well in the lexicon, less in the morphology, phonetics and syntax. According to specialists, up to one third of the Sami lexicon is of substratum origin, and has no analogies in any of the existing languages of the world [20].

Genetic evidence suggests that haplogroup I1,

with which we associate Paleo-European popula-tion of Northern Europe, has the following fre-quencies in the Sami gene pool: Sami – 28% [7], the Sami from Sweden – 32% [22], Inari Sami– 34% [23]*, Skolt Sami – 52% [23], Sami from Lujávri (Lovozero) – 17% [23].

Based on the stated data the hypothesis

about the connection between haplogroup I1 and

Paleolithic population of Europe looks very con-vincing.

Analysis of sources allows us to reconstruct

the population history of haplogroup I1-M253 the following way:

It is not clear yet how and when carriers of

haplogroup I appeared in Europe, this question is being dicussed at present. It is not also clear when and where I1 separated from I. However, in the Paleolithic era carriers of haplogroup I1 settled in in the northern part of central Europe of the territory of present Denmark, northern Germany and Poland. They created such cultures as Ahrensburgian and Swiderian (the author does not exclude the role of carriers of I2a in forming of Swiderian culture); their main occupation was hunting (predominantly for reindeer, elk (Fig. 9) and beaver) and gathering.

Fig. 9. The elk. The skeleton of an elk aged 8700 years was found in a peat bog in 1922 near the town Taderup.

Elk was wounded, in the same bog harpoon was found [26].

_____________________________________________________________ *Haplogroup I1 is not typed in this paper, but the results of other studies [24] and public DNA projects [25] allow us to declare it

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In 11-10th millennium BC global climate changes took place, this led to the melting of the ice cover in Scandinavia. Vegetation penetrated to the territories cleared from the ice cover, with the main food of local population – reindeer – fol-lowing after it; that caused the migration of Pa-leolithic hunters. The colonization of Scandinavia, Baltic and Central Eastern Europe was started.

In the Mesolithic haplogroup I1 faced the

eastern newcomers, who related confidently with haplogroup N1c. They had Uraloid appearance (with a Mongoloid and Caucasoid features). Spreading of Uralic languages in Eastern Europe is connected with N1c.

Relations between the newcomers and abori-

ginal population were generally peaceful; it is evident from mixed graves and gradual appear-ance of mixed anthropological types.

The next important step to the formation of

present situation was made by the carriers of cord ceramics cultures, who had haplogroup R1a [27], and were related to the spreading of South-ern European agricultural and pastoral tribes in Central Europe.

In the 3rd millennium BC tribes of corded ce-

ramics from Central Europe entered the Baltic region (Corded ware culture) and the upper and middle Volga areas (Fatyanovo-Balanovo cul-tures, (Fig. 5)). Their anthropological type was sharply dolichocranic with moderately broad-Caucasoid type [28].

Most likely the tribes of corded ceramics were

quite aggressive and forced out aboriginal popu-lation (I1/N1c) to the remote areas and partially assimilated it. Aboriginal population could not compete with the newcomers economically and

militarily, because it was not familiar with metal-lurgy and productive farming [29].

Mass migration of Slavic tribes (VII-VIII cen-

turies AD) that have made significant changes to the gene pool of Eastern Europe should also be mentioned. The Slavs were mainly the carriers of haplogroup R1a (more) and I2a.

As a result of these processes carriers of Hap-

logroup I1 were partially displaced from their areals and assimilated by more developed new-comers.

In the end I would like to highlight the basic

conclusions: - Roots of haplogroup I1 evidently came from

such Paleolithic cultures as Ahrensburgian and Swiderian; its carriers represented were the part of autochthonous population of Northern and Eastern Europe.

- The main activities of carriers of haplogroup

I1 were hunting and gathering. - Initial anthropological appearance of carriers

of haplogroup I1 was sharply dolichocranic, broad-faced, tall Caucasoid type.

- Carriers of haplogroup I1 were speakers of

Paleo-European language, which didn’t belong to the Uralic or Indo-European families. Its traces were reveiled in the European toponimy and in the Sami language.

Compact local maximum of frequencies of I1

in the center of the Russian Plain is the conse-quence of ancient migrations of Paleolithic popu-lation of Europe, which led to the foundation of Upper Volga culture (the 6-5th millennium BC).

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32. Лобов А.С. Структура генофонда субпопуляций баш-кир. Автореферат на соискание ученой степени канди-дата биологических наук. Уфа. 2009. С.15.

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34. Напольских В.В. Предыстория уральских народов // Acta Ethnographica Hungarica. T. 44:3-4. Budapest, 1999; стр. 431-472.

35. Харьков В.Н., Степанов В.А., Медведева О.Ф. и др. Различия структуры генофондов северных и южных алтайцев по гаплогруппам Y-хромосомы // Генетика. 2007. Т. 43. № 5. С. 675–687.

36. Харьков В.Н., Медведева О.Ф., Лузина Ф.А., Колбаско А.В., Гафаров Н.И., Пузырев В.П., Степанов В.А.. Сравнительная характеристика генофонда теле-утов по данным маркеров Y-хромосомы. Генетика. 2009. Т. 4.

37. Юнусбаев Б.Б.. Популяционно-генетическое исследо-вание народов Дагестана по данным о полиморфизме Y-хромосомы и Alu-инсерций: дис. канд. биол. наук: 03.00.15 Уфа, 2006 107 с. РГБ ОД, 61:07-3/183.

38. Abu-Amero et al Saudi Arabian Y-Chromosome diversity and its relationship with nearby regions // BMC Genetics 2009, 10:59 doi:10.1186/1471-2156-10-59.

39. Battaglia V, Fornarino S, Al-Zahery N, Olivieri A, Pala M, Myres NM, King RJ, Rootsi S, Marjanovic D, Primorac D, Hadziselimovic R, Vidovic S, Drobnic K, Durmishi N, Tor-roni A, Santachiara-Benerecetti AS, Underhill PA, Semino O Y-chromosomal evidence of the cultural diffusion of agriculture in Southeast Europe European Journal of Hu-man Genetics, Vol. 17. No. 6. (June 2009), pp. 820-830.

40. Capelli C, Redhead N, Abernethy JK, Gratrix F, Wilson JF, Moen T, Hervig T, Richards M, Stumpf MP, Underhill PA, Bradshaw P, Shaha A, Thomas MG, Bradman N, Goldstein DB (2003) A Y chromosome census of the British Isles. Curr Biol 13:979–984.

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42. Csányi B., Bogácsi-Szabó E., Tömöry G., Czibula A., Priskin K., Csısz A., Mende B., Langó P., Csete K., Zsolnai A., Conant E.K., Downes C.S. and Raskó I. Ychromosome analysis of ancient Hungarian and two modern Hunga-rian-speaking populations from the Carpathian Basin. Ann Hum Genet 2008. 72: 519-534.

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44. Fornarino S. et al, "Mitochondrial and Y-chromosome di-versity of the Tharus (Nepal): a reservoir of genetic varia-tion," BMC Evolutionary Biology 9:154, 2009.

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45. Heinrich M., Braun T., Sänger T, Saukko P., Lutz-Bonengel S. and Schmidt U. Reduced-volume and low-volume typing of Y-chromosomal SNPs to obtain Finnish Y-chromosomal compound haplotypes Int J Legal Med (2009), 123(5): pp. 413-418.

46. King RJ, Ozcan S, Carter T et al: Differential Y-chromosome Anatolian influences on the Greek and Cre-tan Neolithic. Ann Hum Genet 2008; 72: 205– 214.

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48. Kasperaviciute D, Kucinskas V, Stoneking M. 2004. Y chromosome and mitochondrial DNA variation in Lithua-nians. Ann Hum Genet 68:438–452.

49. Kushniarevich A.I., Sivitskaya L.N., Danilenko N.G. et al. Y-chromosome gene pool of Belarusians – clues from bial-lelic markers study // Proc. Nat. Acad. Sci. 2007. V. 51. № 5. P. 100–105.

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51. Lappalainen et al, Regional differences among the Finns: A Y-chromosomal perspective. Gene Apr 28, 2006.

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55. Marjanovic, D; Fornarino, S, Montagna, S, Primorac, D, Hadziselimovic, R, Vidovic, S, Pojskic, N, Battaglia, V, Achilli, A, Drobnic, K, Andjelinovic, S, Torroni, A, Santa-chiara-Benerecetti, AS, Semino, O (2005). The peopling of modern Bosnia-Herzegovina: Y-chromosome haplo-

groups in the three main ethnic groups. Annals of Human Genetics 69 (6): 757–763.

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57. Noveski et al. Y chromosome single nucleotide polymor-phisms typing by SNaPshot minisequencing // Balkan Journal of Medical Genetics, 12 (2), 2009.

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60. Sengupta S, Zhivotovsky LA, King R et al: Polarity and temporality of high-resolution Y-chromosome distribu-tions in India identify both indigenous and exogenous ex-pansions and reveal minor genetic influence of central Asian pastoralists. Am J Hum Genet 2006; 78: 202– 221.

61. Sengupta S., Zhivotovsky L., King R., Mehdi S.Q., Ed-monds C.A., Chow C-E.T., Lin A., Mitra M., Sil S., Ramesh A., Usha Rani M.V., Thakur C.M., Cavalli-Sforza L., Ma-jumder P., Underhill P. «Polarity and Temporality of High-Resolution Y-Chromosome Distributions in India Identify Both Indigenous and Exogenous Expansions and Reveal Minor Genetic Influ-ence of Central Asian Pastoralists»// Am. J. Hum. Genet. 78 (2): 202–21.

62. Varzari A. (2006), "Population History of the Dniester-Carpathians: Evidence from Alu Insertion and Y-Chromosome Polymorphisms," Dissertation for the Facul-ty of Biology at Ludwig-Maximilians University, München.

63. Zastera J., Roewer L., Willuweit S., Sekerka P., Benesova L., Minarik M. Assembly of a large Y-STR haplotype data-base for the Czech population and investigation of its substructure , 13 July 2009 Forensic Science Internation-al: Genetics April 2010 (Vol. 4, Issue 3, Pages e75-e78).

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Arabian clusters of haplogroup E1b1b1c1 (M34)

A.A. Aliev, D.L. Tartakovsky

Abstract

Haplogroup E1b1b1c1* (M34) and its subclade E1b1b1c1a* (M84) were detected among the Arabs in the Ara-bian Peninsula. A possible reason for migration of the founder of cluster E1b1b1c1a-E from the Levant to the Ara-bian Peninsula could be the Crusades.

Theme talk

The highest diversity of subclades of hap-logroup E1b1b1c1 (M34) is observed in the Levant and Anatolia, therefore its ancestral home is often placed in the Eastern Mediterra-nean [1, 2, 3]. In addition, haplogroup E1b1b1c1* (M34) and its subclade E1b1b1c1a* (M84) were detected among the Arabs in the Arabian Peninsula [4, 5], where they form spe-cific clusters — E1b1b1c1-B [6] and E1b1b1c1a-E [7]. Knowing the age of the clusters and their area of distribution, we can find out the history of clusters’ origin and resettlement of their carriers. In this paper we will try to find out the history of E1b1b1c1 and E1b1b1c1a subclades in the Arabian Peninsula on the ex-ample of these clusters.

Arabian clusters: when and why? To find out the origin of the clusters, let us

define their ages with the probability of 95% ac-cording to [8]. At the time of writing the paper (July 2010) cluster E1b1b1c1-B has had only two 67-marker haplotypes (N=2). Obviously, due to such a small number of haplotypes, their TMRCA (time to most recent ancestor) is “too young” and is 350±320 years, and gives us no reason to draw any definite conclusion about the history of its origin.

The sample of cluster E1b1b1c1a*-E con-sists of five 67-marker haplotypes. This clus-ter’s TMRCA is 1090±510 years.

Despite the fact that, due to different size of

samples, the ages of these clusters’ founders are different, it should examine the entire period of their confidence intervals, which are inter-sected. It is possible that both clusters have ari-sen in about the same medieval era and are linked to the migration of their founders from the Levant to the Arabian Peninsula according to some important event. What could cause this mi-gration?

We think that a possible cause of the mediev-

al migrations from the Levant could be the Cru-sades — a series of Western invasions to oust the Muslims from Palestine, which lasted almost two hundred years (1096 - 1272 years).

The first crusade ended with the capture of

Jerusalem and the massacre of Muslims [9]. Apparently, these invasions, and, as a re-

sult of them, looting and killings, forced part of the Muslims to seek refuge from persecu-tion of the Crusaders closer to Mecca. This, in our view, could cause to arise at least one cluster of Arabia — E1b1b1c1a-E.

_____________________________________________________________

Received: July 16 2010; accepted: July 18 2010; published: July 19 2010 Correspondence: [email protected]

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Conclusions 1) Carriers of subclades E1b1b1c1* (M34)

and E1b1b1c1a (M84) identified in the Ara-bian Peninsula, where they form clusters E1b1b1c1-B and E1b1b1c1a-E.

2) The TMRCA of cluster E1b1b1c1*-B is

50±320 years ago, the TMRCA of cluster

E1b1b1c1a*-E is 1090±510 years ago. They pos-sibly arose at one time.

3) A possible reason for migration of the

founder of cluster E1b1b1c1a-E from the Le-vant to the Arabian Peninsula could be the Cru-sades.

References

1. C. Cinnioğlu et al. (2003), «Excavating Y-chromosome hap-

lotype strata in Anatolia». Hum Genet (2004) 114 : 127-148. DOI 10.1007/s00439-003-1031-4

2. Mirvat El-Sibai, Daniel E. Platt, Marc Haber, Yali Xue, Sonia C. Youhanna, R. Spencer Wells, Hassan Izaabel, May F. Sanyoura, Haidar Harmanani, Maziar Ashrafian A. Bonab, Jaafar Behbehani, Fuad Hashwa, Chris Tyler-Smith, Pierre A. Zalloua. Geographical Structure of the Y-chromosomal Genetic Landscape of the Levant: a coastal-inland con-trast. Annals of human genetics, 2009.

3. A. A. Aliev, Bob Del Turco. Modern carriers of haplogroup E1b1b1c1 (M34) are the descendants of the ancient Le-vantines. Russian Journal of Genetic Genealogy. Vol 1, 2010.

4. Haplozone E3b, Arabian E-Y-DNA Project, Arab DNA Project.

5. Cadenas et al. (2007), «Y-chromosome diversity characte-rizes the Gulf of Oman», European Journal of Human Ge-netics 16: 1–13, doi:10.1038/sj.ejhg.5201934

6. E1b1b1c1*-B cluster 7. E1b1b1c1a*-E cluster 8. Адамов Д. Расчет возраста общего предка по мужской

линии для «чайников». The Russian Journal of Genetic Genealogy (Русская версия), Том 2, №1, 2010 г.

9. Раймунд Ажильский, История франков, которые взяли Иерусалим (Raimundi de Aguiliers. Historia Francorum qui ceperunt Iherusalem) в кн. «История крестовых по-ходов в документах и материалах», М., 1975 г.

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Origin, Distribution and Migrations of I2b*-Subclades (18 september 2008 – http://sites.google.com/site/haplogroupil38/)

Hans De Beule

Abstract

Until now the resolution of most scientific articles was not detailed enough to say something about the small

haplogroup I2b*. The purpose of this paper is to describe the continental origin, distribution and migrations of the I2b*-subclades. To calculate a minimum spanning network 101 European I2b* samples were used. Starting from this network, clusters within the known I2b* subclades (I2b*-A, -B and –C) were determined by combining DYS448 and DYS19 values.

Origin of the samples, distribution and place of origin of the surname were taken into account to pinpoint the Continental samples (together with the related British Isles samples) on the map of Europe. The Upper Rhine re-gion clearly played a prominent role in the history of I2b*. This region has the highest frequency of I2b*s and the greatest cluster-diversity.

Introduction

I2b* (old I1b2*; positive for SNP’s: S23, S30, S32, S33; negative for M223) is an old and ro-bust clade that originated about 4500 years ago in northern Europe. I2b* consists out of three subclades –A, -B and –C. According to Ken Nordtvedts’ (2008) modal values spreadsheet for haplogroup I; I2b*-A is characterized by DYS448 = 19, I2b*-B by DYS448 = 21 and I2b*-C by DYS448 = 20.

I2b* can be linked to 3000 to 2700 years old

skeletons found in the Lichtenstein cave in the German Harz mountains. Thirteen of the 19 male skeletons found there, can be determined as be-longing to haplogroup Ib2*. Culturally these founds belong to the Unstrut-culture (between the river Unstrut and the Southern Harz moun-tains) which:

is rooted in the Funnelbeaker Culture (also

called TRB or Trichterbecher) – 4000 BC to 2700 BC - characterized by gatherer-hunters becoming farmers;

is seen as one of the cultures that lead to

the first Germanic culture: the Jastorf culture – 6th to 1st century BC.

Because of the known distribution of the re-

lated M223 clade I2b1 ( I1b2a or I1c in the old naming conventions), it appears that the point of origin of I2b* should be looked for in the valleys of the river Elbe.

Figure 1: distribution of I1c (current I2b1) as presented by Wiik (2008).

_________________________________

*Remark: In 2008 I-L38 (aka I2b2 in 2010) was known as I2b*.

_____________________________________________________________

Received: July 28 2010; accepted: July 30 2010; published: August 5 2010 Correspondence: [email protected]

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Seen this information it is tempting to con-clude that I2b* is linked to the Albe and to the TRB-culture. The results of this study however do not match this theory.

Subjects

In order to explain the distribution of I2b* in Europe 5 approaches were combined:

1. calculation of the minimum spanning net-

work; 2. clustering of the samples in subclades; 3. studying of the historical origins of the

samples within each cluster; 4. studying of the distribution and first place

of occurrence of the sample’s surnames; 5. calculation of the MRCA between Conti-

nental en British Isles-samples within each clus-ter.

Methods

1. Calculation of the minimum spanning network and clustering of the samples

According to Bandelt (1999) the multitude of

plausible phylogenies trees is best expressed by a network which displays alternative potential evolutionary paths. A minimum spanning tree for a set of sequence types connects all given types, such that the total length (the sum of distances between linked sequence types) is minimal. The minimum spanning network serves as a good point of departure to reconstruct the most likely tree by taking geographical information into ac-count. The Median Joining Networks in this paper are created by the Fluxus 4.5 Software.

To create a minimum spanning network for

I2b*, 10 STR values for 98 samples were used, all selected out of the ysearch (2008) and SMGF (2008) databases. All samples have the following values: DYS454 = 12, DYS455 = 10 and DYS448 =19, 20 or 21.

As an historical point of reference the values for 3 Lichtenstein-individuals Y1, Y2 and Y3 were added (although the DYS448 values for these samples were predicted).

The STR-loci used were: DYS 19, 392, 389i,

448, 389ii, 385a, 385b, 391, 439, 390. Loci with identical values for all samples were not used. Appendix A refers to the used samples which STR-values are online available.

Number and origin of the 101 I2b* samples

1 Belgium (BEL) 6 France (FRA) 2 Netherlands (NET) 12 Scotland (SCO) 2 Switzerland (SWI) 13 Germany (GER) 3 Denmark (DEN) 53 England (ENG) 6 Ireland (IRE) 3 samples out of the

Lichtenstein cave (Y1, Y2 and Y6)

From different samples with identical sur-

names only the sample with the most known markers was used. Non-European I2b* samples were not used because it proved too difficult to pinpoint them to a European location of origin. As an exception only a few non-European samples with a known European origin were included.

In the Fluxus software it is optional for the

entered STR-values to choose between a stan-dard weight (of 10) or a customized weight (with a maximum weight of 100).

For the STR-values, weights based on muta-

tion rates (Chandler 2006) were entered. For ex-ample: locus DYS19 has a mutation rate of 0.00151 per generation -or- 1 mutation every 662,25 generations; this means 1 mutation every 16 556 years (662,25 x 25 years per generation). Because of the mutation rate of 1/16k years, six-teen was entered as a weight.

2. Clustering of the samples in subclades

To determine to which I2b* subclade the

samples of the network core belong, Jim Cullen’s Haplo-I-Subclade Predictor was used (see Figure 2). This predictor works on a weighted genetic distance algorithm. In basic terms, the predictor makes a large number of random sample obser-

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vations of the entered haplotype and predicts, for each observation, which modal haplotype best describes the sample of markers. Each modal haplotype is rated in percent by its ability to best describe the sample of markers during the trials. The Haplo-I-Subclade Predictor is based on sub-clade modal values and geographic distributions from the research of Ken Nordtvedt.

3. Studying the historical origins of the samples within each cluster

In some cases the ysearch database links STR-values to a name, birth date, date of death, place of birth/death of the most distant known paternal ancestor of the sample. Although not all ysearch users choose to post this information on the website, it often contains valuable informa-tion. This information should be seen as indica-tive, since it is very difficult to check. In some cases, when the origin of the sample was not en-tirely clear, the family was contacted and asked for additional information.

4. Studying the origin and first place of occurrence of a sample’s surnames

Surname research sometimes gives a hint of the likely cultural background of a sample. In some cases the information is highly indicative, in others most speculative. As an additional histori-cal source surname research is meaningful. For an overview of the origins of I2b* surnames see Appendix B.

As Bowden (2007) argues, the link between

surname and Y-chromosomal haplotype is imper-fect, due to multiple founders for names and his-torical non-paternities and adoptions.

Nevertheless unrelated men sharing surnames

are significantly more likely to share haplotypes than are men carrying different names. This demonstrates that surnames have been associ-ated with specific haplotypes for many genera-tions and suggests that access to the Y-chromosomal diversity of past populations might be possible through the selection of modern samples based on surnames known to exist in a particular region during the medieval period.

Bowden (2006) describes that to maximize the benefits of surname-based ascertainment, care needs to be taken in sampling. Common surnames should be avoided where possible. The more frequent names are likely to have had mul-tiple founders and may provide less reliable links to a specific region.

For all Continental samples and all samples

mentioned in the MRCA paragraph below, the dis-tribution of the surname was mapped (see Fig-ures 7 and 8) when:

1. the sample was not already linked to a

known location; 2. the name did not rank among the most

popular surnames. For example: the distribution of the surnames of the three Danish samples was not taken into account since they refer to the 3rd, 4th and 5th most popular Danish surnames);

3. the distribution map had a clear geo-

graphic centre. In case there were two centres, the location of first occurrence of the surname was selected.

5. Calculation of the MRCA between Continental and British Isles-samples within each cluster

To calculate the MRCA between Continental and British Isles haplotypes the maximum of available STR-information was used. Mostly, 37 markers (the first 3 FTDNA panels) were used. When this was not possible, 32 or 25 markers were used to calculate the MRCA.

FTDNA’s population geneticists state that 25

years best expresses a typical generation prior to the Dark Ages (476-1000 AD) and 25 to 30 years per generation for the period thereafter. Since this paper covers both timeframes MRCAs were calculated for generations of 25 years as well as for generations of 30 years.

To calculate the genetic distance between 2

haplotypes, each single point mutation was counted as a mutational event.

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Example of a MRCA calculation with a genetic distance of «10»:

The average mutation rate of the 37 markers

of the first three FTDNA panels is 0.004952. This means the average mutation rate for the 37 markers is 1/202 per generation.

Taking into account 37 markers this means

one mutation every 5.5 generations (202/37). Calculating with generations of 25 years this means one mutation every 136 years. A genetic distance of 10 equals 1360 years between two haplotypes -or- 1360/2 = 680 years between both involved haplotypes and their MRCA.

Results Calculation of the minimum spanning network

Figure 2 shows the core of the minimal span-ning network with DYS448 subclades indicated. Figure 3 displays the entire network. As the col-our codes in the networks show, the combination of the values of DYS19 and DYS448 clusters the samples within a subclade.

Nodes representing several samples: BEL1 also represents NET2; DEN2 also represents DEN3, ENG7,

ENG44, ENG45, ENG46, ENG47, ENG48; SCO5 also represents SCO6;

ENG14 also representing ENG41;

ENG26 also represents ENG29; ENG31 also represents ENG36;

ENG39 also represents ENG40, ENG42; FRA6 also represents ENG4;

SCO3 also represents SCO4;

SCO10 also represents ENG25;

Y1 also represents GER13;

Y6 also represents IRE4, IRE6, ENG38,

ENG43.

The phylogenetic network displays the rela-tionship between the inserted weighted STR-values and shows all possible evolutionary trees. This means that the original Fluxus network dis-played much more links between the nodes. To eliminate the unlikely links, thus to reconstruct the most likely tree, all available information was used:

comparison with networks with alternative

settings; close inspection of STR-values of the sam-

ples of the network’s core; genetic distance;

available geographic information;

haplogroup prediction (Cullen, 2008).

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Figure 2: I2b* subclades of the network core. Nodes represent haplotypes and are proportional to the number of sampled individuals. The codename of the sample refers to the country of origin of the sample. The length of the links represents

the genetic distance. The colour of the nodes refers to specific DYS19 and DYS448 combinations.

Figure 3: the most likely I2b* network displaying all 101 samples.

Hg I prediction: ENG53 I2b*-C: 49% I2b*-B: 39%

Hg I prediction: ENG31 I2b*-A: 36% I2b*-C: 23% ENG36 I2b*-C: 49% I2b*-B: 32%

Hg I prediction: Y2 I2b*-C: 94% I2b*-A: 3%

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Clustering of the samples

The table and pie charts below summarize the subclade frequencies for:

all samples; the Continental samples; the British Isles samples.

I2b*-

C I2b*-B ? I2b*-A

Sum Cont. 1* 7 2 2 3 1 3 8 4 31 Isles 7 19 14 0 1 5 2 14 8 70 Total 8 26 16 2 4 6 5 22 12 101

* = predicted

Of the 101 samples (including the 3 Lichten-stein samples) 47% belongs to I2b*-B, 39% to I2b*-A, 8% to I2b*-C and 6% to the undefined orange cluster (with DYS19=16 and DYS448=20). Due to the overrepresentation of British Isles samples the overall pie chart is probably not representative.

8%

25%

16%

2%4%

6%

5%

22%

12%

Figure 4: I2b* frequencies per subclade.

Looking at the 31 Continental samples, 45%

belongs to I2b*-B, 49% to I2b*-A and only 3% (in casu the Lichtenstein sample with the pre-

dicted DYS448 value) belongs to I2b*-C and 3% to the undefined orange cluster.

Figure 5: Continental I2b* frequencies per subclade.

Of the 70 British Isles samples 49% belongs

to I2b*-B, 34% to I2b*-A, 10% to I2b*-C and 7% to the undefined orange cluster.

Figure 6: British Isles I2b* frequencies per subclade.

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Studying the historic origin of the samples within each cluster

For details or an overview of the known geo-graphic origin of the samples, please refer to ap-pendix A. All small circles, displayed on Figure 7 below, refer to samples with a documented geo-graphic place of origin.

Studying the origin and first place of occurrence of the sample’s surnames

Concerning the surname distribution several (free and online available) sources have been used:

the English surname distribution was

based on the England and Wales census records of 1891;

the Scottish surname distribution was

based on the 1891 Scotland census records;

the Irish surname distribution was based on the Primary Valuation property survey of 1848-1864 (per household);

the French surname distribution was

based on the census of 1891-1915;

the Dutch surname distribution was based on the phonebook-entries of 1993;

the Belgian surname distribution was

based on the census of 1998;

the German surname distribution was based on phonebook-entries of 2002.

In all cases the known information of the

samples, the surname distribution and the loca-tion of first occurrence of the surname was used as a check before pinpointing a location with a small triangle on the map below.

The little square refers to one case where the

surname itself refers to a Dutch locality.

Figure 7: the distribution of Continental and related British Isles I2b* samples. Circles represent known geographic origins of the samples, triangles represent the areas with the highest frequency of a sample’s surname, a square represents

the origin of a locational surname.

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Calculation of the MRCA between Continental en British Isles samples within each cluster

To learn more about the historic relation of

the Continental and British Isles samples, the MRCAs were calculated between Continental samples and the samples linked to it, whenever:

the Continental sample has at least three

direct links to other samples; the related samples belong to the same

DYS448-DYS19 cluster as the Continental sam-ple.

The Lichtenstein samples were not taken into

account. The age estimates of common ancestry of

Continental and British Isles samples should be

interpreted with caution. One should keep in mind that the MCRA age estimate does not reflect an actual crossing of the Channel. We can as-sume that the crossing of the Channel took not place before the MRCA date. This date limits the historic scenarios.

For example: the MRCA of a German and Eng-

lish sample could have lived between 1145 and 1289 AD, while the forefathers of the English sample crossed the channel as French Huguenots in the 16th century. In this case we know only that the Channel was not crossed before 1145-1289 AD.

Figure 8 displays the geographic relationship

between Continental and British Isles samples that are linked in the I2b*-network (see Figure 3).

Continental Sample Related with …

The MRCA calculated with

generations of 30 years lived

around …

The MRCA calculated with

generations of 25 years lived

around …

DEN2 Tryk ENG49 Hutchinson 578 BC 147 BC SCO5 Cruikshank 362 BC 33 AD SCO6 Harris (adopted)

362 AD 33 AD

NET2 Lems (BEL1 De Beule)

ENG12 Wootan 316 BC 71 AD IRE1 Finley 150 BC 209 AD ENG22 Brooks 182 AD 486 AD

GER10 Zimmer ENG10 Mortimer 348 AD 625 AD ENG5 Cockrell 779 AD 983 AD GER2 Seiler (Say-lor)

942 AD 1120 AD

GER4 Wehr FRA1 Brion 205 AD 506 AD SCO12 Garscadden 679 AD 901 AD IRE2 Holland 499 AD 751 AD ENG1 Boucher 779 AD 983 AD FRA6 Le Roi 861 AD 1052 AD ENG13 Bassett 861 AD 1052 AD

GER13 Underwood ENG2 Chapman 715 AD 930 AD ENG43 Holmes 930 AD 1110 AD ENG34 Clark 930 AD 1110 AD IRE4 Brabazon 1145 AD 1289 AD IRE6 Bellew 1145 AD 1289 AD ENG38 Sawyer 1361 AD 1469 AD

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Figure 8: relations between the Continental and British Isles I2b* samples (cfr. the MCRA table above).

Discussion

Analysing the I2b*-network supports the clus-tering of I2b* in 3 major subgroups (I2b*-A, -B and –C) by using DYS448. It also shows that DYS19 (=DYS394) can be used to identify haplo-type-clusters within these subclades.

Analysis of the network shows that the com-

binations of DYS19 and DYS448 are not at ran-dom. These clusters seem to refer to distinct mi-gration waves.

In fact the DYS19, DYS448 combination might

reveal how the I2b* tree evolved. Looking at the network route (see Figure 3) between the green (I2b*-A) and the white (I2b*-B) clusters, DYS19, DYS448 evolves from 15,19 to 16,19 to 16,20 to 16,21 and finally to 17,21. To determine which direction of this network route is upstream and which downstream further research is necessary to determine the age of I2b*-A, -B and –C.

Migration scenario per cluster I2b*-A

Most continental I2b*-A’s are found in the Upper Rhine region (for a definition or map of the Upper Rhine, please refer to http://en.wikipedia.org). The Upper Rhine is the location with the highest number of continental samples and the greatest Ib2*-diversity. This seems to be the historic starting point of several I2b*-A subclades.

The Upper Rhine region corresponds to the

territories of the Germanic Istvaeones as men-tioned by Tacitus (55-118AD).

Green cluster hypothesis (DYS19=15, DYS448=19)

This I2b*-C subcluster seems to have mi-

grated to the Low Countries at an early point in time. The MRCA calculations suggests that they crossed the Channel before the 4th – 5th century. The structure of the network (see the connection

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of BEL1, NET1, NET2 and GER7 which is con-nected to an Irish sample with the surname «Hol-land») and the MRCA calculations supports an Upper Rhine to Low Countries to British Isles mi-gration. Orange cluster hypothesis (DYS19=16, DYS448=20)

The DYS448 and DYS19 values of the orange

cluster seems to reflect an intermediary step be-tween I2b*-B and I2b*-C. The only direct conti-nental link is sample GER9 which comes from the Upper Rhine region.

Yellow cluster hypothesis (DYS19=16, DYS448=19)

The Upper Rhine sample GER4 of the yellow cluster seems to be the absolute centre of I2b*-A. MRCA calculations suggest that from this point a migration wave to the British Isles started around the 11th century. The samples and sur-names involved seem to point to a strong relation with the Norman invasion.

I2b*-B

The I2b*-B cluster has several centres: The white cluster (DYS17=15,

DYS448=21) starts in Denmark and links the continent (Denmark) to Yorkshire. The MRCA cal-culation indicates a migration in between the 3th of 4th century). This might correspond with a mi-gration of Anglii or Cimbrii.

The red cluster (DYS19=16, DYS448=21)

seems to connect North Germany to Yorkshire and Ireland. Looking at the MRCA between the continental sample GER13 and the connected British Isles samples it looks as if the migration took place during/after the battle of Hastings (1066 AD).

Another red cluster centre seems to be

connected to the main I2b* centre in the Upper Rhine region. Here too, the connected British Isles-samples suggest a connection to the Nor-man-invasion of the Isles.

I2b*-C

The samples of the blue I2b*-C cluster (DYS19=15, DYS448=20) have no direct links to the continental samples; although the predicted haplogroup of Lichtenstein sampleY2 points in the direction of this haplogroup.

Probably this cluster was a Germanic group

that left for the British Isles at a very early stage. The limited amount of DYS19=15, DYS448=20) samples are located in North-England, Ireland and Scotland.

Oppenheimer’s (2006) genetic analysis shows

that there were major Scandinavian incursions into northern and eastern Britain during the Neo-lithic period and before the Romans.

General conclusion

On the European continent, the frequency of I2b* as well as the diversity of DYS448 and DYS19 combinations is highest in the Upper Rhine region.

The concentration of the continental I2b*’s in

the Upper Rhine region, along the Rhine, needs further examination.

Is this the location of origin of I2b* (or some

of its subclades) or did I2b* (or some of its sub-clades) migrate to this location? If the former is true it might help to explain why the distribution of I2b* is mysteriously low in North Europe.

In both cases the question is which people

and cultures were involved? Is there a relation to the LBK-finds (Linear Pottery Culture) in the Up-per Rhine region? Or do we need to look for Celtic or Germanic migrations?

In order to solve this gigantic puzzle it would

be very revealing to look for other (sub)haplogroups that correlate with the I2b*-subclades and compare their distributions.

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Webreferences

1. Free network software: http://www.fluxus-engineering.com 2. Haplogroup I subclade modals: http://knordtvedt.home.bresnan.net/FounderHaps.xls 3. Haplogroup I predictor: http://members.bex.net/jtcullen515/haplotest.htm 4. STR databases: http://www.smgf.org http://www.ysearch.org/ 5. Surname distribution maps: English and Scottish surnames: http://www.ancestry.co.uk/facts Irish surnames: http://irishtimes.com/ancestor/surname German surnames:

http://christoph.stoepel.net/geogen/v3/Default.aspx

French surnames: http://www.geopatronyme.com Belgian surnames: http://www.familienaam.be/ Dutch surnames: http://www.familienaam.nl/ 6. Explanation of surnames: German surnames: http://www.duden.de/duden-

suche/werke/famnamen/ French surnames: http://www.jeantosti.com/noms/t1.htm English and Scottishsurnames:

http://www.ancestry.co.uk/facts and http://www.houseofnames.com/

References 1. Bandelt Hans-Jürgen, Forster Peter, Röhl Arne.(1999) Me-

dian-Joining Networks for Inferring Intraspecific Phylog-enies. Molecular Biology & Evolution, 16(1): 37-48.

2. Bowden Georgina R., Balaresque Patricia, King Turi E., Hansen Ziff, Lee Andrew C., Pergl-Wilson Giles, Hurley Emma, Roberts Stephen J., Waite Patrick, Jesch Judith, Jones Abigail L., Thomas Mark G., Harding Stephen E., Jobling Mark A. (2008) Excavating Past Population Struc-tures by Surname-Based Sampling: The Genetic Legacy of the Vikings in Northwest England. Molecular Biology and Evoution. 25(2):301–309.

3. Capelli Cristian, Redhead,Nicola, Abernethy Julia K., Gra-trix Fiona, Wilson James F., Moen Torolf, Hervig Tor,Richards Martin, Stumpf Michael P.H., Underhill Peter A., Bradshaw Paul, Shaha Alom, Thomas Mark G., Brad-man Neal, Goldstein David B. (2003) A Y Chromosome Census of the British Isles. Current Biology, Vol. 13, 979–984, May 27.

4. Chandler John F. (2006) Estimating Per-Locus Mutation Rates. Journal of Genetic Genealogy, 2:27-33.

5. Schilz Felix (2006) Molekulargenetische Verwandtschafts-analysen am prähistorischen Skelettkollektiv der Lich-tensteinhöhle. Dissertation, Göttingen.

6. Tacitus. Germania. Ambo-Klassiek, 1992, pp. 175-205. 7. Oppenheimer Stephen (2006) Myths of British Ances-

try.Prospect Magazine. Issue 127, October 2006. 8. Weale Michael E., Weiss Deborah A., Jager Rolf F., Brad-

man Neil, Thomas Mark G. (2002). 9. Y Chromosome Evidence for Anglo-Saxon Mass Migration.

Molecular Biology and. Evoution. 19(7):1008–1021. 10. Wiik Kalevi. 2008. Where did European Men Come From?

Journal of Genetic Genealogy, 4:35-85.

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Appendix A: Samples Network

ID User ID

Surname Origin sample

Y1, Y2, Y6 Lichtenstein cave

750 BC, Osterode am Harz, Germany

GER1 P88QG Strohmeier 1649-1729 Bogen, Bavaria, Germany

GER2 U9HMG Saylor Jacob Seiler, 1715-1793

GER3 4BRM9 Ochs °1625 Poppenhausen Germany

GER4 RQS47 Wehr °1720 Heidelberg, Pfalz, Germany

GER 5 66196 Krassin 1791 - 1798 Kolmar, Posen, Poland

GER6 SMGF Tietjen

GER7 6WNX5 Steinmetz °1756 Germany

GER8 VUERB Hartung 1620-1700 Geisleden, Germany

GER9 9TD9J Marschall °1755 Ommeray, Lorraine, Germany

GER10 BBB59 Zimmer °1866 Darmstadt, Germany

GER11 8VWK5 Creswick Schmidt 1820-1868, Upper Rhine,Lorraine, Germany

GER12 X22KV Greene °1790 Germany

GER13 X8EDM Underwood 1832-1865 Berlin

DEN1 SMGF Hansen °1742 Denmark

DEN2 TWRDQ Tryk

DEN3 N9812 Peder Andersen 1742, Hojrup, Tonder

BEL1 N14392 De Beule °1560, Zele, Belgium

NET1 SMGF Van Hoesen

NET2 SMGF Lems 1504 Rotterdam, Netherlands

FRA1 8GD73 Brion Kirrberg, Elzas, France

FRA2 JHH9G Rogers Tancred de Hauteville 975-1078, France

FRA3 58EBF Guittard

FRA5 SMGF Tavernier

FRA6 50508 Le Roi

SCO1 3G37R Findley °1795 Scotland

SCO2 CC6CC Hutchison

SCO3 6D9UQ McKinney

SCO4 PJ7UT MacLeay

SCO5 F5M64 Harris-adopted

SCO6 UN3VU Cruikshank

SCO7 FFKC9 Adam

SCO8 NAJ27 Parks

SCO9 X5F8D McClellan

SCO10 2AADH McKinzey

SCO11 BTE2U Levack

SCO12 SMGF Garscadden

IRE1 JYWUE Finley °980 Ireland

IRE2 AWMBB Holland °1780 Ireland

IRE3 K3V8G Menary °1841 N-Ireland

IRE4 BPKEY Brabazon 1692-1772, Ballin-voher, French Park,IRE

IRE5 F9J8G Walden 1841, N-Ireland

IRE6 GHSTC Bellew 1710-1776 Mount-bellew Co Galway, Ireland

ENG1 BAYSF Boucher

ENG2 GY8X3 Chapman

ENG3 9GDC6 Doane

ENG4 BQ4UU Lay

ENG5 BKVDK Cockrell °1807 England

ENG6 8PF4W Phillips

ENG7 5DZHE Berry

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ENG8 HDNJD Terry

ENG9 8GEWA Daniel °1765 England

ENG10 ex56

2YGGC Mortimer °1635 -1704 Wilt-shire England

ENG11 VBT8X Morrel

ENG12 7CBQN Wootan °1620 England

ENG13 NAZCH Bassett °1830, Llanelli, Wales, England

ENG14 NYYDS Pittman

ENG15 R3EY7 Chewning Chowning, 1620-1660 Wotham, Kent, England

ENG16 WXSVN Butler 1819-1905, Co-lerne, Wiltshire, England

ENG17 J7JWB Scharschmidt

ENG18 BKAY5 Mills

ENG19 P2894 Weakley 1695-1743, Mar-tock, England

ENG20 SMGF Ellis

ENG21 FG6NF Rust

ENG22 B7J2M Brooks °1690, Lancaster-shire, England

ENG23 DXF2E Cullen 1579, Upton by Southwell, Notting-hamshire, England

ENG24 SMGF Milner

ENG25 VX6H2 Oldfield Hall, 1813-186 Hanley castle, Wor-cestershire, England

ENG26 A996E Weathers °1696 England

ENG27 F6NNW Miller

ENG28 QHMNK Evans 1854-1920 Ha-worth, Yorkshire, England

ENG29 WFS7K Withers °1695 England

ENG30 ZVCW3 Campbell

ENG31 EEVS5 Hamblin °1588 England

ENG32 GFEDE Moses

ENG33 6JQN9 Todd 1620 England

ENG34 6CXZ5 Clark

ENG35 SZ9FV Foster 1595 England

ENG36 9U6JZ Furbey °1840-1892 Whit-nash, Warwickshire, England

ENG37 UGUGG Greenwood

ENG38 4ZSBU Sawyer 1623-1702 Bed-fordshire, England

ENG39 584DC Speak °1698 England

ENG40 JCE48 Speake

ENG41 RAYMJ Rawls °1745 England

ENG42 RPGYZ Payne

ENG43 XMAJP Holmes Holme, 1632-1703 England

ENG44 WQUH5 Stanley

ENG45 DZ53W Brinley

ENG46 SMGF Worthington

ENG47 SMGF Bennett

ENG48 VNQYP Weston

ENG49 E77WQ Hutchinson 1779-1838 Aldby, England

ENG50 QP5ZD Dodd

ENG51 GFCJX Flory

ENG52 Z9X3R Gilmore

ENG53 PEU8S Riviere The name Fox was changed to Riviere in 1895.

SWI1 MN9NA Lehman °1702 –1778, Schauffausen ,Switserland

SWI2 7F3ME Flora Fleury, 1682-1741 Switzerland

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Appendix B: About the Network’s Surnames

GER1: Strohmeier: This sample comes from Bogen, Bavaria, Germany.

GER2: the name Saylor is a variant of Seiler. GER3: Ochs: Low-German surname, meaning

ox. Low German was spoken in Westphalia. The Ochs sample is traceable to 1625 in Poppen-hausen, Germany. The surname itself was first found in the Rhineland (Westphalia).

GER4: Wehr: this surname refers to the local-

ity Wehr in Reinland-Pfalz, Germany. The sample is traceable up to 1720 in Heidelberg, Pfalz. To-day most German Wehr’s are found in the landkreis Eichsfeld.

GER5: Krassin: his sample is traceable until

1791 in Kolmar, Posen, Poland (ancient Prusia). GER6: Tietjen: today most Tietjens live in

North Germany in the Landkreis Osterholz. GER7: Steinmetz: this German sample is

traceable until 1756. This occupational name has its highest concentration in the landkreis Trier – Saarburg.

GER8: Hartung: this sample is traceable until

1620-1700 in Geisleden, Germany. GER9: Marschall this sample is traceable until

1755 in Ommeray, Lorraine (département Moselle), France.

GER10: Zimmer; this sample is traceable until

1866 in Darmstadt, Germany. The highest con-centrations of German families with the surname Zimmer is found in the German landkreis Saar-louis.

GER11: Creswick: the Schmidt sample is

trackable til 1820-1868 in Upper Rhine/Lorraine Germany.

GER12: Greene GER13: Underwood; the sample is traceable

til 1832 in Berlin; the surname is traceable until 1791 in Kolmar, Posen, Poland (old Prusia).

DEN1: Hansen: Danish patronym: son of Hans.

DEN2: Tryk: Danish surname traceable until

1742 in Tønder nearby Branderup. BEL1: De Beule: this sample is traceable until

1560 near Dendermonde, Flanders, Belgium. NET1: Van Hoesen: this surname refers to the

Dutch locality Huizen in North-Holland. First found in Haarlem in 1388 (Baertout van Huesen).

NET2: Lems: this family has a traceable fam-

ily tree up to 1504, in the neighbourhood of Rot-terdam at the mouth of the Rhine. Most Lems to-day live in south-west Netherlands, along the North sea coast.

FRA1: Brion: this sample is traceable up to

Kirrberg, Upper Rhine, France. The name is common in entire France.

FRA2: the Rogers sample is traceable in direct

paternal line to Tancred de Hauteville (975-1058) in France. This Norman was a minor noble in Normandy.

FRA3 Guittard: common in Puy-de-Dôme and

in the region Tarn, surname of Germanic origin, Widhard (wid = wood + hard = hard).

FRA4: Long: frenck for tall. In France this

surname is common in the southeast. This name is also widespread in Great-Britain.

FRA5: Tavernier: French occupational name

common in Picardie and Nord-Pas-de-Calais. FRA6: Le Roi: French for the king. This sur-

name is common in the Nord-Pas-de-Calais and in Picardie. The related surname Le Roy is com-mon in Bretagne and in Normandy.

SCO1: Findley: from the Scottish name

Fionnlagh / Fionnlaoich, meaning «fair hero» - from the Gaelic elements «fionn» meaning white or fair and «laoch» meaning warrior or hero. First found in Banfshire in the northeasterly Grampian region of Scotland, where they were descended

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from the chiefs of the Clan Farquharson, one of the great clans, known as the clan Chattan.

SCO2: Hutchison First found in Northumber-

land were they were seated from very early times, some say well before the Norman con-quest (1066).

SCO3: MacKinney: Irish names, first found in

the Irish county Monaghan, where they were they were known as the lords of Truagh.

SCO4: Mac Leay: Gaelic name (Origin Gaelic)

The son of Clay.

SCO5: Cruikshank Scottish surname. First found in Kincardineshire where they held a family seat from very ancient times.

SCO6: Harris (adopted) SCO7: Adam: The surname Adam is of great

antiquity in Scotland. Duncan Adam, son of Alex-ander Adam, lived in the reign of King Robert Bruce (1274-1329), and had four sons, from whom all the Adams, Adamsons, and Adies in Scotland are descended.

SCO8: Parks: English and Scottish: from Mid-

dle English, Old French parc a metonymic occu-pational name for someone employed in a park or a topographic name for someone who lived in or near a park. In the Middle Ages a park was a large enclosed area where the landowner could hunt game.

SCO9 McClellan SCO10: McKinzey: variation of SCO3. SCO11: Levack: The Levack name appears in

Caithness records from about the mid 1600s. It is said to be affiliated to the MacLea and the Living-stons.

SCO12: Garscadden: Scottish name with lots

of spelling variations. First found in Dumbarton-shire(Gaelic: Siorrachd Dhn Bhreatainn), pres-ently the Council Areas of West and East Dunbar-tonshire, where they were anciently seated, some say before the 12th century.

IRE1: Finley Scottish: from the Gaelic per-sonal name Fionnlagh (Old Irish Findlaech), com-posed of the elements fionn ‘white’, ‘fair’ + laoch ‘warrior’, ‘hero’, which seems to have been rein-forced by an Old Norse personal name composed of the elements finn ‘Finn’ + leikr ‘fight’, ‘battle’, ‘hero’.

This sample recently comes from Dublin, Ire-

land, but much much earlier from Balchristie, Fife, Scotland, and before that the west coast of Scotland, and before that the east cost of Ire-land... and before that... we get deep into the myths of Macbeth's father's lineage

son - Macbeth Fionnladh b.c. 1005 Atholl,

Perthshire, Scotland; son - MacBeatha McFinlay b. 1045 Cromarty,

Ross and Cromarty, Scotland; d. 1093 Cromarty, Ross and Cromarty, Scotland; md. 1079 Bethoca McBrad daughter of Andrew McBrad;

son - Ruari (Rory) McFinlay b. 1080 Cromarty,

Ross Cromarty, Scotland; son - Fergus McFinlay b. 1145 Aberdeenshire,

Scotland; son - Eugenius McFinlay b. 1184 Perthshire,

Scotland; son - Fearchar McFinlay b. 1210

son - Archibald Finlay b. 1248 Roushknot, Perthshire, Scotland;

son - William Finlay b. 1300 Perthshire, Scot-

land; son - Andrew Finlay b. 1344 Perthshire, Scot-

land; son - John Finlay b. 1390 Perthshire, Scot-

land; son - John Finley b. 1418 Perthshire, Scot-

land; son - John Findley b. 1450 Coupar Angus,

Perthshire, Scotland;

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son - Andrew Finley (Fyndlay) b.c. 1480/1483 of Perthshire, Scotland;

son - James Finley b. Sept. 15, 1530 Cuper

Angus, Balchristie, Fife, Scotland. IRE2: Holland: this Irish surname, refers to

the Netherlands -or- is a reduced Anglicized form of Gaelic Ó hÓileáin, a variant of Ó hAoláin, from a form of Faolán (with loss of the initial F-), a personal name representing a diminutive of faol ‘wolf’.

IRE3: Menary: Irish surname with lots of

variations and a possible french Huguenot origin. IRE4: Brabazon: this surname refers to

someone from the duchy of Brabant (Belgium). By the thirteenth century, it was also an occupa-tional name for a mercenary, specifically a mem-ber of one of the more or less independent ma-rauding bands of mercenaries, noted for their lawlessness and cruelty, who originated in Bra-bant but in the course of time accepted recruits from almost anywhere The earliest of the name recorded was Tomas Brabazon, listed as a tenant in the Domesday Book of 1086. Other records of the name mention Thomas Brabezon, 1273, in Yorkshire county.

IRE5: Walden: habitational name from any of

the places, in Essex, Hertfordshire, and North Yorkshire, named Walden, from Old English w(e)alh ‘foreigner’, ‘Briton’, ‘serf’+ denu ‘valley’.

IRE6: Bellew: first found in Yorkshire were

they were granted lands by William the Con-queror after the Norman conquest in 1066. This family came to Ireland around 1200 with the Normans from England and settled in the East in Counties Meath & Louth and then to the west in Co. Galway in the 1650's.

ENG1: Boucher (Origin French) A butcher; a

blood-thirsty man. French and English: occupa-tional name for a butcher or slaughterer, Middle English bo(u)cher, Old French bouchier (also with the transferred sense ‘executioner’), a derivative of bouc ‘ram’. This sample is traceable until 1740 in Virginia where they were part of the Scots-Irish migration wave. The theory is that they were French Huguenots who fled to Ireland

where they mixed with the Scots who also mi-grated to Ireland.

ENG2: Chapman: the same as Chipman, a

trader, a shopman; from the Saxon ceapan or cypan, to buy or sell.

ENG3: Doane: Anglo-saxon topographic name

for a downland dweller (from Old English dun ‘down’, ‘low hill’), first found in Cheshire were they were seated from very early times, some say well before the Norman conquest.

ENG4: Lay: variant of Lee. ENG5: Cock(e)rell: Middle English for cock-

erel, a young cock. ENG6 Philips: patronymic from the personal

name Philip. ENG7: Berry: from the province of Berri, in

France. First found in Devonshire, where they were granted lands by William the Conqueror af-ter 1066.

ENG8: Terry English and Irish: from the

common Norman personal name, T(h)erry (Old French Thierri), composed of the unattested Germanic element þeudo- ‘people’, ‘race’ + ric ‘power’. Theodoric was the name of the Os-trogothic leader (c. 454–526) who invaded Italy in 488 and established his capital at Ravenna in 493. His name was often taken as a derivative of Greek Theodoros. There was an Anglo-Norman family of this name in County Cork.

ENG9: Daniel: from the Hebrew personal

name Daniel ‘God is my judge’. ENG10 Mortimer Norman name. First found in

Herefordshire, where there were seated from early times and were granted lands by William of Normandy, their liege lord, for their assistance at the battle of Hastings in 1066 AD. This sample was traceable until 1635 in Wiltshire.

ENG11: Morrel: Having yellow hair. First

found in Norfolk where they were seated from early times and were granted lands by Duke Wil-liam of Normandy after their assistance at the battle of Hastings 1066.

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ENG12: Wootan: English: habitational name from any of the extremely numerous places named with Old English wudu ‘wood’ + tun ‘en-closure’, ‘settlement’. This name is related to the Wootten, Woten, Wooten and Wooton families. This anglo-saxon name was first found in Kent were they were anciently seated at Marley before and after the Norman conquest. Today Wootan is numerous in Kent and Lancashire.

ENG13: Welch sample with a surname derived

from the French Basset: a little fat man with short legs and thighs.

ENG14: Pittman English: topographic name

for someone who lived in a hollow –or- German (Pittmann): probably from a compound personal name formed with Pitt, a short form of Peter + Mann ‘man’.

ENG15: Chewning: variation of the name

Chew: which refers to the Anglo-Saxon personal name Ceawa. First found in Somerset were the family were granted lands by William of Nor-mandy for their assistance at the battle of Hast-ings 1066 A.D.

ENG16: Butler: this family derives their origin

from the old Counts of Briony or Biony, in Nor-mandy, a descendant of whom, Herveius Fitz Walter, accompanied the Conqueror into England. His son, Theobold, went with Henry II. into Ire-land, where, having greatly assisted in the reduc-tion of the kingdom, he was rewarded with large possessions there. The king afterward conferred on him the office of chief Butler of Ireland.

ENG17: Scharschmidt: although an English

sample, a German name. ENG18 Mills: Scottish and English: topog-

raphic name for someone who lived near a mill. ENG19: Weakley: variant of Weekley? habita-

tional name from a place in Northamptonshire called Weekley, from Old English wic ‘settlement’, perhaps in this case a Roman settlement, Latin vicus + leah ‘wood’, ‘clearing’.

ENG20 Ellis: Contracted from Elias.

ENG21: Rust English (chiefly East Anglia) and Scottish: nickname for someone with red hair or a ruddy complexion, from Old English rust ‘rust’ (from a Germanic root meaning ‘red’). First found in Kent were they were anciently seated as lords of the manor.

ENG22 Brooks; derivation of «brook», or a

small stream. Also a name given to those who came from Brooksbank, the name of several places in England. First found in Essex where they were granted lands by William the Con-queror for their assistance at the battle of Hast-ings.

ENG23: Cullen: habitational name from the

Rhineland city of Cologne. When the name ar-rived into England shortly after the events of 1066, the name was changed from «de Cologne» to various spellings, most commonly as Cullen, a popular variant at the time. The known male line of this sample extends back to the early 1500's in Nottinghamshire, England.

ENG24: Milner Northern English (mainly York-

shire) and Scottish: variant of Miller, retaining the -n- of the Middle English word, which was a result of Scandinavian linguistic influence, as in Old Norse mylnari.

ENG25: Oldfield: This name is of Anglo-Saxon

origin, and is locational from any of the various places thus called: Oldfield in Yorkshire, Worces-tershire or Cheshire. The surname Oldefeld was first recorded in 1297 in Yorkshire.

ENG26: Weathers/Withers: AngloSaxon

names, first found in Hampshire where they were seated from very early times before and after the Norman invasions.

ENG27 Miller: English and Scottish: occupa-

tional name for a miller. ENG28: Evans:Welsh for John. First found in

Herefordshire where they were seated from very ancient times (before the Norman conquest).

ENG29: variation of ENG26. ENG30: Campbell: the origin of the name can

be Scottish, Celtic or Gaelic. The ancient Camp-

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bell family may be traced as far back as the be-ginning of the fifth century in Lochore, Ar-gyleshire, Scotland?

ENG31: the surname Hamblin is a corruption

of Hammeline, which was taken from Hamelen, a town on the river Weser, Germany. In England the surname was first found in Gloucestershire where they were seated from very early times and were granted lands by William of Normandy for their assistance at the battle of Hastings 1066 AD.

ENG32: the surname Moses has endless spell-

ing variations (Moy, Moye, Moyes, Moesen, Moi, …) which is characteristic for a Norman surname. The name was first found in Shropshire and they were anciently seated as lords of the manor.

ENG33: Todd: scotch name for a fox, first

found in Berwickshire were they were seated from early times.

ENG34: Clark, a clergyman, a scholar, one

who can read and write. ENG35: Foster: English: reduced form of

Forster. ENG36: this name refers to the little Norse-

Viking village of Fearby in North Yorkshire. The sample is traceable til 1840 in Whitnash is War-wickshire.

ENG37: Greenwood:English: topographic

name for someone who lived in a dense forest, from Middle English grene ‘green’ + wode ‘wood’, or a habitational name from a minor place so named.

ENG38: the surname Sawyer was first re-

corded in Norfolk England where they were seated from early times and their first records appeared on early census rolls taken by the early kings of Britain to determine the rate of taxation of their subjects.

ENG39: English: nickname for someone

thought to resemble a woodpecker in some way, Middle English spek(e) (a reduced form of Old French espeche(e), of Germanic origin).

ENG40 Speake: cfr Speak ENG41: Rawls: patronymic from a medieval

form of the personal name Ralph. First found in Cornwell were they were anciently seated as Lords of a manor.

ENG42 Payne: from the Latin Paganus, now

out of use, meaning a man exempt from military service.

ENG43: Holmes: English (mainly Yorkshire)

and Scottish: topographic name for someone who lived by a holly tree, from Middle English holm, a divergent development of Old English hole(g)n; the main development was towards modern Eng-lish holly.

ENG44: Stanley: a market-town in Glouces-

tershire, England. The place of a tin mine, stan, tin, Welsh, ystaen, and ley; or from the Saxon, stan, a stone, and ley--the stony place.

ENG45: Brinley: first found in Cheshire were

they were seated from very early times, some say well before the Norman conquest (1066).

ENG46: Worthington: first found in Lanca-

shire, before and after the Norman conquest. The name Worthington is derived from the locality whence the family came. Its etymology is three Saxon words. 'Wreath in ton' that is. 'Farm in town'. Twenty miles north-east of Liverpool in Leyland hundred, parish of Standish, county of Lancaster, England is the town Worthington. Here and in the adjacent manors resided the family of Worthington for many generations, being estab-lished, from the time of the Plantagenets (who ruled the Dutchy of Normandy in between 1144-1204).

ENG47: Bennett: first found in Lancashire;

Dutch, Scottish or English origin. ENG48: Weston: first found in Staffordshire

having be granted lands as a tenant in chief by William the Conqueror.

ENG49: Hutchinson: the son of Hitchins or

Hutchins (Hugh).

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ENG50 Dodd: from the Middle English per-sonal name Dodde, Dudde, Old English Dodda, Dudda, which remained in fairly widespread and frequent use in England until the 14th century. It seems to have been originally a byname, but the meaning is not clear; it may come from a Ger-manic root used to describe something round and lumpish—hence a short, plump man.

ENG51: Flory: cfr SWI2. ENG52: Gilmore: Gaelic, the henchman or fol-

lower of the chief, one who carried the chief's broadsword, from gille, a servant, and mor, large, great.

ENG53: Riviere: in 1895 a Fox adopted his mother’s surname Riviere (French for river).

SWI1 Lehman: this sample was traceable til 1702 in Schaufhausen, Switzerland.

SWI2: Flora / ENG51: Flory: The name of

Flory/Flora in Germany was often spelled Flori, a spelling that has close associations with Switzer-land. In Britain the name was first found in Som-erset.

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Origins of Hg I-L38 (I2b2) Subclades (5th of April 2009 - http://sites.google.com/site/haplogroupil38/)

Hans De Beule

Abstract

Network analysis confirms I-L38-B (and especially the Lichtenstein variant) as the ancestral

I-L38 clade. Distribution of continental I-L38 samples with known geographical origin confirms the Upper Rhine area as region with the highest I-L38 frequency and diversity. Distribution of I-L38 (I2b2) in the Netherlands does not support a Saxon ancestry.

Further analysis is needed to clear out the relation of I-L38 to the Rhine, to study the possible link to early bronze age cultures as the Michelberg culture and to investigate the correlation to other clades as R-L21*.

Introduction

As conversations on [email protected] pointed out I-L38 is thinly spread over Europe; from Italy and Spain to Slo-venia, Switzerland, Germany, Denmark, the Netherlands, Belgium and the British Isles. I-L38 is almost absent in Scandinavia and East Europe. As Ken Nordtvedt remarked: this spread indicates an old haplogroup, or possibly a hap-logroup travelling faster than others. The distribution of I-L38 does not correlate that of haplogroup I-P30 (I1a). I-L38 seems to have leaked into France, as if it arrived before I-P30.

A previous paper (De Beule, 2008) pointed

out that on the European continent, the fre-quency of I-L38 as well as the diversity (demon-strated by various DYS448 and DYS19 combina-tions) is highest in the Upper-Rhine region (see Figure1). This contradicts the supposed origin of I-L38 in the middle Elbe region. This view is mainly built on the finds of the I-L38 Lichtenstein bones in the Harz mountains and the high fre-quency of I-M223 (I2b1) in this region. I-L38 and I-M223 however separated already during the LGM. There is no reason to assume a correlation between I-L38 and I-M223.

Distribution of Haplogroup I-L38 in the Netherlands

Recently Barjesteh van Waalwijk van Doorn published «Zonen van Adam in Nederland» (suns of Adam in the Netherlands). This book presents DNA-profiles of 410 Dutch ‘suns of Adam’ and describes the distribution of haplogroups in the Netherlands. The 410 samples were linked to ge-nealogical information and geographical location of the oldest known ancestor in the male line.

The haplotyping was done by the FLDO (Fo-

rensic Laboratory for DNA research) based in Lei-den, Netherlands. Of the 410 samples, 8 be-longed to I-L38 (belonging to 5 families – see Table 1 below).

The haplotyping was done by the Haplogroup

I predictor and was based on 16 markers: DYS 393; 390; 19; 391; 385a; 385b; 439; 389i; 392; 389ii; 458; 437; 448; H4; 456; 438.

It is remarkable that almost all pinpointed

I-L38 samples can be found very near main riv-ers; in casu the Rhine, Neckar, Scheldt and Meuse.

_____________________________________________________________

Received: July 28 2010; accepted: July 30 2010; published: August 5 2010 Correspondence: [email protected]

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Figure 1: distribution of Continental and related British Isles I-L38 samples. Dots represent known origins of samples, triangles and squares are educated guesses based on the sample’s surname. The colours indicate specific DYS19-DYS448 clusters

as a demonstration of I-L38 diversity (De Beule, 2008).

Table 1: Dutch I-L38-families of the «Zonen van Adam in Nederland» study.

Surname Hg I prediction

I-S23 refers to I2b2

Location of the oldest known

forefather Remarks on the geographic locations

Blaas I-S23-A =>36% I-S23-A-RecLOH =>36% I-S23-C =>16% I-S23-B =>8%

Rees (Germany) Near the river Rhine.

De Booij I-S23-B =>48% I-S23-A =>24% I-S23-A-RecLOH =>24% I-S23-C =>5%

Kleve (Germany) Near the river Rhine.

Lems I-S23-C =>60% I-S23-A =>19% I-S23-A-RecLOH =>19% I-S23-B =>1%

Hoogvliet (Netherlands)

This haplotype is similar to that of the Belgian De Beule family, with an histori-cal origin around Zele along the river Scheldt (near St Amands) Hoogvliet is located near of the mouth of the Rhine.

Meert I-S23-B =>96% I-S23-A =>2%

St Amands (Belgium)

Village near the river Scheldt (also very near Zele).

Spée I-S23-A =>48% I-S23-A-RecLOH =>48% I-S23-B =>2%

Baarlo (Netherlands)

Near the river Meuse.

Lichtenstein cave Osterode am Harz middle Elbe region

UpperRhine region

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Whereas the coastal distribution of hap-logroup I (dominantly I-M253 and I-M223) in the Netherlands supports a Saxon origin; the distri-bution of I-L38 does not. If I-L38 had a Saxon origin the samples would be found grouped to-gether with the other haplogroup I samples along the Northsea coast and in Frisia (see Figure 2).

Network of Continental I-L38

This network analysis is performed as de-scribed in the paper «Origin, Distribution and mi-grations of I2b*»-paper. The network below is based on 33 continental I-L38 samples with known geographical background coming from a number of sources (see Appendix B).

The sample (#33) is small; but highly indica-

tive since I-L38 is a small haplogroup. Whenever the samples were included in «Ori-

gin, Distribution and migrations of I2b*», the same code name is used. This is why the sample code not seems coherent at first sight.

Figure 2: distribution of haplogroup I in the Netherlands - map based on locations of origin of the oldest known

forefather – blue dots represent I-L38.

Continental subclades I-L38-A and I-L38b-B

Network analysis (see Figure 3) supports the

view that haplogroup I-L38 basically consists out of two continental varieties: I-L38-A and I-L38-B. Hopefully a SNP will be found soon to determine I-L38 subclades. In the meantime it is pragmatic to distinguish I-L38-A versus I-L38-B.

Figure 3: continental I-L38 network with a red line dividing I-L38-A and I-L38-B. The colours refer to the DYS19-DYS448 clusters as described in «Origin, Distribution and migrations of I2b*subclades».

I-L38-A

I-L38-B

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The network also suggests that subclades I-L38-D, I-S23-A-RecLOH and I-L38-15 could be seen as varieties of I-L38-A and I-L38-14 could be a variety of I-L38-B. I-L38-C seems to be a Scottish variant of I-L38-B.

Instead of identifying subclades or clusters,

this paper categorizes the samples either in hap-logroup I-L38-A or I-L38-B. This is done based on STR-values, the Haplogroup I Predictor and the network results. In the network, the Lichtenstein node Y1 (also representing sample GER13 from Berlin) seems to be the root of all other branches. Distribution of Continental I-L38

Based on the network structure it is tempting

to relate the geographical origin of I-L38 to the

geographical location of the Lichtenstein cave. Since I-L38 samples are found from the source to the mouth of the Rhine it probably makes more sense to see the spread of continental I-L38 in relation to the Rhine.

There is a high and divers concentration of

I-L38 in the Upper Rhine region: As a previous paper (De Beule, 2008) indi-

cated there is a high concentration of I-L38b-A in the Upper Rhine region. Also see Fig-ure 1.

As a paper in preparation by Steve Ralls on I-

L38b-14 shows; 35% of all known (#17) I-L38-14 samples comes from the Upper Rhine region (35% is related to the British Isles and the re-maining 30% is spread from Sweden to Italy).

Figure 4: the distribution of Continental I-L38 samples based on the known origin of 33 samples. Red dots representing I-L38-B, yellow dots I-L38-A. The callout marks the location of the Michelberg, the green arrows displays a hypothetical migration route

of the ancestral I-L38.

I-L38 map

Michelsberg

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Discussion and Further Investigation Migration hypothesis

Starting from the Upper Rhine, some groups probably followed the Rhine downstream (possi-bly even crossed the Channel), others followed the Rhine downstream. The ancestral I-L38 vari-ety seems to have travelled along the Rhine til Kleef / Rees and from there on travelled east to the area of the Lichtenstein cave in the Harz mountains. See Figure 4.

To explain these migrations they should be

seen in their historical context (i.e. in the context of the emerging bronze age). The Harz region (where the Lichtenstein cave lies) was an early centre of copper/bronze and the Rhine was an evident trade route for bronze objects.

Michelberg culture

The distance of both I-L38-A and I-L38-B to their Most Common Recent Ancestor is 135 gen-erations (calculated with Ken Nordtvedt’s «Gen-erations» spreadsheet). Calculated with genera-tions of 30 years this means their MCRA lived 4800 years ago. With a standard deviation of 25.7% this implies a MCRA living between 5850 and 3750 years ago. In Germany this timeframe refers to the transition of the Late Neolithic into the Early Bronze Age.

Looking at the distribution of artefacts belong-

ing to the Michelberg culture a pattern emerges that resembles the continental distribution of I-L38 (see Figure 5 compared to Figure 4).

The Michelsberg culture blossomed from ca.

6400 to 5500 years ago. For more information on the Michelberg cul-

ture, see Appendix A. It is interesting that this culture links the Upper Rhine region to the Middle Elbe.

Figure 5: distribution of the Michelberg culture. Correlation with R-L21*

The spread of I-L38 also resembles the spread

of R-L21* (R1b1b2a1b6*) – see Figure 6. Just as I-L38 (see Figures 1 and 4), haplogroup R-L21* seems connected to the course of the Rhine and to the British Isles.

Conclusion

The deeper one digs into the history of I-L38 the more relations appear with the Upper Rhine area and bronze age cultures.

The separation of I-L38-B and I-L38-A seems

to have taken place in the early bronze age in Germany. Looking at the distribution of I-L38, rivers (and especially the Rhine) seem to have played an important role. It is along these rivers that bronze objects were distributed.

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Figure 6: the distribution of R-L21*(cfr. the R-L21 Project).

May be the different clusters of I-L38 can lead to a better understanding of the historical migra-tions up and down the Rhine. May be there even was a relation to copper/bronze that could ex-plain the I-L38 presence in the Harz mountains (a known prehistoric centre of copper and bronze)

and the British Isles. In this respect it also is in-teresting to look at the cultural sphere of the Michelberg culture. May be there even is a rela-tion to R-L21* which seems to have a similar dis-tribution than I-L38.

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Webreferences

1. Haplogroup I predictor: http://members.bex.net/jtcullen515/haplotest.htm 2. Ken Nordtvedts Generations Spreadsheet http://knordtvedt.home.bresnan.net/Generations2.xls 3. DNA-list focussed on haplogroup I: http://archiver.rootsweb.ancestry.com/th/index/Y-DNA-

HAPLOGROUP-I/

4. FTDNA L38 project: http://www.familytreedna.com/public/I2b2/default.aspx 5. I2b2 project website: http://tinyurl.com/2b76jd 6. R-L21 project website: http://www.familytreedna.com/public/R-L21/default.aspx

References

1. Hans De Beule. Origins, distribution and migrations of

I2b*subclades posted on dna-forums.org on 18 Septem-ber 2008 (in 2008 I-L38 was still called I2b*).

People interested in this paper can find it at the FTDNA I-L38 project site (see URL above) or can email me.

2. Steve Ralls. Paper on I-L38-14 (in preparation).

Appeal

To learn more about I2b2 it is crucial to be able to pinpoint a sample to a geographical loca-tion. This is why the group administrator of the FTDNA L-38 project Tim Weakley urged all I2b2’s

who joined the project , to enter in the informa-tion (name and location if possible) for their most distant known ancestor in the male line. Please do!

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Appendix A: Michelberg culture

For more information on the Michelberg cul-ture, refer to: http://www.comp-archaeology.org/Michelsberg.htm

A few interesting quotes: The Michelsberg culture is named after the

Michelsberg, a hill at Untergrombach, Kr. Bruch-saal, Baden-Württemberg, Germany and llasted from ca. 4400-3500 cal BC. The Michelsberg sites distribution includes the area around the Middle Rhein (Rhine) River, Belgium and the Paris Basin. A few sites with Michelsberg pottery are reported from Central Germany and the Czech Republic.

In Germany, Schumacher saw parallels be-

tween the Michelsberg culture and the Late Meso-lithic pottery making Kjøkkenmødinger culture (now Ertebølle culture) as early as 1908 (Lüning 1969). Similarities with the Funnel Beaker culture (TRB) pottery were also noted and some re-searchers still include the Michelsberg culture in the TRB interaction sphere (i.e. «the TRB in the larger sense».

Until the 1960’s the culture was seen as part

of the «lake dwelling sphere of the northern Alps and its piedmont», which includes Aichbühl, Pfyn and Horgen. In the 1960’s the Michelsberg cul-ture was separated from these cultures and seen as evolving out of Bischeim, a late phase of the Rössen culture in the Middle Rhein (Rhine) River region of Germany.

On the other hand, the American archaeolog-

ist Scollar (1959, 1961) stressed the Michelsberg culture’s western origin. Similarly Dubouloz

(1998) argued that the pottery of the Paris Basin was possibly older than in Germany and sug-gested that its development should be associated with the Menneville Group of «Early Post-Rössen» and the initial phase of the Early Chasséen (Chassey) culture (Jeunesse 1998).

In Belgium the pottery assemblage of Spiere

«de Hel» began to be seen as a kind of western development that exhibit a geographically transi-tional subgroup, resembling Michelsberg and to some extent the neighboring cultures (Vanmont-fort 2001, Vanmontfort et al. 1997).

In the east (Central Germany, Bohemia

and possibly even Moravia) Michelsberg-like pottery occurs during the Baalberge Phase of the TRB’s Middle-Elbe and South Group, which begins to form around 4000/3800 cal BC.

In the South Group and adjacent regions the

Baalberge Phase gives way to TRB Phase II sometime between 3600/3400 BC (Baldia et al. in press a, in press b, exhibiting similarities to the Boleráz Phase of the Baden culture. Baden «influence» is even suggested on the Central German Schöninger Group (Raletzel-Fabian and Furholt 2006). In southern Germany near the Bodensee (Lake Constance), the Michelsberg cul-ture gives way to the Horgen culture around 3600/3400 cal. BC. In the Northwest German state of Hessen (Hesse) and adjoining regions Michelsberg evolves into the Wartberg culture at that time. At the same time Michelsberg is re-placed by the later Funnel Beaker Culture (TRB or Middle Neolithic I) at its northernmost fringes.

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Appendix B: samples

User ID Network ID Surname Origin sample

Y1 Osterode am Harz Y2 Osterode am Harz Y6 Osterode am Harz P88QG GER1 Strohmeier 1649-1729 Bogen, Bavaria U9HMG GER2 Saylor 1715 Methingen Metzingen 4BRM9 GER3 Ochs °1625 Poppenhausen RQS47 GER4 Wehr °1720 Heidelberg, Pfalz 66196 GER5 Krassin 1791 - 1798 Kreis, Kolmar, Posen VUERB GER8 Hartung 1620-1700 Geisleden 9TD9J GER9 Marschall °1755 Ommeray Lorraine BBB59 GER10 Zimmer °1866 Darmstadt 8VWK5 GER11 Creswick b1820 Fredrich Schmidt Upper Rhine Lorraine X8EDM GER13 Underwood 1832 Berlin 2MCE9 GER14 Schlenke 1861 Bosseborn (Kreis Hoxter)

Ancestry GER15 Heltzel Johann Tobias Heltzel [also Höltzel] , b. 1732 in Palatinate, Germany, d. 23 November 1792, Paradise, York Co., Pa.

FTDNA 105008

GER16 Roland Gasper Roland, b. ca. 1721, Palatine, Germany, died ca. 1709, Drake Creek, Warren Co., Ky.

ZvAiN GER17 Blaas Rees Germany

ZvAiN GER18 De Booy Kleve, Germany N14392 BEL1 De Beule 1560 Zele ZvAiN BEL2 Meert St Amands, Belgium SMGF NET2 Lems NET, Rotterdam, 1504

ZvAiN NET3 Spée Baarlo, Netherlands 8GD73 FRA1 Brion Kirrberg, Elzas, FRA N25287 FRA7 Hauteville-la-Guichard, France 53868 FRA8 Jean GUITTARD Bellemagny, Alsace

140263 FRA9 Claude REYNAUD

Bâtie Montgascon(38),France

N17917 NOR1 Ommund Ommundson

Fjellestad,b.1812 Norway.

E2623 POL1 Adalbertus/Wojciech Tatucha 1750

Warta,Lodz,Poland

MN9NA SWI1 Lehman °1702 Schauffausen SWIT 7F3ME SWI2 Flora 1682 Joseph Jacob Fleury was listed as a French

Huguenot from Palatinate Germany, though an-other report lists Solothurn, Switzerland

W3MJW SLO1 Wanchick Jastrabie, Slovakia SMGF ITA1 Gandola Primo Gandola, b. ca. 1812, Bellagio, Italy N9812 DEN3 Peder Andersen 1742, Hojrup, Tonder

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Early Bronze Age Origin and Late Iron Age (La Tène) Migrations of I-L38 (November 2009 - http://sites.google.com/site/haplogroupil38/)

Hans De Beule

Abstract

I-L38 is a small clade with a continental distribution scattered around the Upper Rhine (Rhineland - Palatinate).

It also is present on the British Isles. This paper tries to reveal how I-L38 migrated from its continental core to the British Isles. In order to do so, 3

methods have been combined: Firstly, a trendline was calculated for the geographical Y and X co-ordinates of samples with known origin

(belonging to I-L38 and other haplogroups). Secondly, a phylogenetic tree was made for I-L38 samples with known origin. Thirdly, the historical context was studied. Combining these approaches lead to the conclusion that: - starting from the Upper Rhine, I-L38 spread during the EBA in an area between Rhine, Danube and Elbe; - I-L38 migrated in the Late Iron Age I-L38 with Celtic La Tène people, through Belgium, to the British Isles.

Introduction

Haplogroup I-L38 is defined by the SNPs L38/S154, L39/S155, L40/S156, L65/S159. In the ISOGG tree its current name is haplogroup I2b2. It is an ancient clade with a limited number of members.

1. Main clusters of I-L38

Until now, no SNP was found to separate I-L38 (although L39 looks promising).. Several researchers structured I-L38 using different markers into different clusters. Initially Ken Nordtvedt separated I-L38 into 3 clusters using DYS448:

I-L38A with DYS448=19; I-L38B with DYS448=21;

I-L38C with DYS448=20.

Network analysis (De Beule September 2009)

visualized the separation between I-L38A (DYS448=19) and I-L38B (DYS448=21). In this network some of the DYS448=20 samples are positioned as intermediary nodes between I-L38A and I-L38B.

To be consistent with previous papers, this

paper mainly focuses on I-L38A and I-L38B. In this approach I-L38-14 can be considered

as a cluster within I-L38B and I-L38D as a cluster within I-L38A.

2. The EBA separation of I-L38 Using the Generation Spreadsheet of Ken

Nordtvedt it seems that I-L38A and I-L38B sepa-rated 135 generations ago (error range: +/-35 generations). Calculated with generations of 31

_____________________________________________________________

Received: July 28 2010; accepted: July 30 2010; published: August 5 2010 Correspondence: [email protected]

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years this means that the two clades separated 4185 years ago; this is during the EBA (Early Bronze Age) which started in Germany 4400 to 4200 years ago. This estimate of age is useful since from that moment on it becomes possible to track and compare the migrations of multiple clusters of I-L38.

3. The Upper Rhine origin of I-L38 As De Beule (September 2008 and April 2009)

pointed out the distribution of continental I-L38 samples with known geographical origin confirms the Upper Rhine area (Rhineland – Pa-latinate) as region with the highest continental I-L38 frequency and diversity; thus as the likely point of origin of I-L38. For a map of the Rhine see: http://en.wikipedia.org/wiki/File:Rhein-Karte.png

4. Virtual absence of I-L38 in the Netherlands

Integration of data of the Dutch project

De zonen van Adam in Nederland lead to the conclusion that I-L38 is virtually absent in the Netherlands. This implies that historical scenarios that involve north Germanic people (Jutes, Fri-sians, Angles, Saxons, …) became unlikely.

Although there are very few Belgian data

available, two Belgian I-L38 samples were lo-cated around the river Scheldt. Ongoing research http://www.brabant-dna.org/joomla/ will improve the knowledge on Belgian DNA.

This is important since Belgium is geographi-

cally positioned on the most likely route from the Upper Rhine to the British Isles. The distribution of I-L38 in the Low Countries (Belgium, Nether-lands, Luxemburg) seems to be related to the rivers Rhine, Meuse and Scheldt. (De Beule Sep-tember 2008).

5. Re-evaluation of the Michelberg scenario

Based on the distribution De Beule (April 2009) suggested a link between the Michelberg culture and I-L38. This, however, does not fit the

MCRA calculation mentioned in Method 2. Michel-berg is dated from 4200 to 3500/3400 BC and at best could be linked to ancestors of the I-L38 MCRA. Michelberg culture also does not explain the relation between the Upper Rhine region and the British Isles.

6. The EBA Upper Rhine connection

Connecting the Upper Rhine region to the EBA it is interesting that the first EBA groups in the Rhine, Danube and Elbe/Saale regions (Adler-berg, Singen, Straubing, Unetice) appeared around 2400/2200 BC. (Mail dr. Dirk Fabian).

Traditionally, the EBA in Southern Germany has been subdivided into several separate groups such as Adlerberg, Singen, Straubing, Neckar, Upper Rhine, etc on the basis of grave goods and funeral practices. Upon closer examination this separation appeared questionable. Instead, it be-came apparent that within the EBA there are more elements, which are common to these groups than differences. (Libber, 2004) In other words: there must have been interaction (gene exchange) between these groups.

Figure 1: Spread of EBA groups in southern German (Libber 2004 - after Kraus 1988).

7. I-L38 and the Lichtenstein cave

The Lichtenstein cave (Schilz 2006) links I-L38 to the (pre-)Urnfieldculture (1300-700 BC). In this cave, in Osterode-am-Harz 3000 year old bones were found. Y-DNA analysis categorized them as I-L38 (# 13), R-S21 (# 1) and R1a (# 2).

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Recently, two direct descendants of the I-L38 bones were found, living in the valley next to the cave, proving that haplotypes can stay at one location during millennia. The known STR-values of the Lichtenstein bones are shown in Appendix 1.

Inserting these STR-values in Jim Cullen’s

Haplogroup I Predictor the most likely hap-logroups are:

- Haplotype Y1: I-S23-A: 41%; - Haplotype Y2: I-S23-C: 72%;

- Haplotype Y4: I-S23-A: 33%;

- Haplotype Y6: I-S23-B: 67%;

- Haplotype Y3: R1b-Frisian: 32%;

- Haplotype Y5: R1a: 33%.

Looking at the I-L38 predictions it is remark-

able that all three DYS448 clusters seem to be present in the Lichtenstein cave.

8. I-L38 and related haplogroups

The diversity of haplogroups in the Lichten-stein cave confirms that during the Bronze Age populations consisted out of several haplogroups. So, haplogroups whose distribution is related to the distribution of I-L38 might learn us some-thing about I-L38. In a previous paper Hans De Beule suggested that the distribution of some R1b subclades seemed to mirror that of I-L38.

In this context it is interesting to mention pro-

fessor Steve Jones, a Welsh geneticist. In an arti-cle in Y Faner Newydd he linked R1b to the Celtic culture and separated the most important R1b clades into the following scheme:

- S21/U106 (River Celts) – common in Aus-

tria particularly around the western core Urnfield-Hallstatt area, along the Rhine to the Netherlands and down the Danube to Bulgaria. This haplogroup was also found in the Lichten-stein cave.

- S28/U152 (Alpine Celts) – common in the Alps in regions of Alpine Germany, Switzerland and Northern Italy, but also from Greece to the Bay of Biscay.

- L21/S145 (Insular Celts) – common in the

Celtic Isles of the North-West coast of Europe such as Britain, Ireland, the Isle of Man, etc, but also found in France, Germany and Scandinavia.

- M153, M167 (Iberian Celts) – common in

regions of Spain and Portugal with a Celtic-Basque-Iberian heritage such as Minho, Galicia, Asturias, Cantabria, Euskara, Catalonia and down to Andalusia, but also in the Celtic Isles, France and Germany. (http://www.celticheritage.org/SteveJones.php)

The next paragraph tries to clarify the possi-ble relations between I-L38 and R1b-clades with a method based on geographical co-ordinates.

Method 1: geographical analysis

To investigate the relation between I-L38 and other subclades, it is interesting to focus on the possible relation with the R1b «River», «Alpine» and «Insular Celts». It also is worthwhile to see whether there is a geographic relation between I-L38 and its brother clade I-M223.

To investigate the geographical relation be-

tween I-L38 and the subclades mentioned above, locations of the oldest known ancestor were se-lected for each clade from FTDNA projects. Sometimes the available data needed to be re-duced (e.g. for R-S1 every 10th row was used. If this sample did not display geographical informa-tion, the next sample in line was used, etc).

Only for I-L38, extra locations were added

from the Dutch research De zonen van Adam in Nederland.

For the selected data the decimal geographi-

cal co-ordinates were looked up. For each subclade, these co-ordinates were

inserted in an Excel spreadsheet. All inserted data were displayed in a graph and for each sub-clade Excel was used to calculate a trendline.

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Using the trendlines: - it is possible to compare the trendlines

(although the result should be treated with cau-tion);

- it is possible to compare the trend to

known historical locations/points of reference; As points of reference geographical coordinates of the following historical locations were added to the graph: EBA-locations (Adlerberg, Singen, Straubing, Neckar, Upper Rhine, Unetice); Celtic locations (Hallstatt, La Tène) and the Lichtenstein cave (in Osterode-am-Harz).

- it is possible to translate points of interest

(eg. two crossing trendlines) to geographical co-ordinates. Conclusions (figure 2):

Starting from the Upper Rhine region - The trendlines of I-L38A and I-L38B to

the west are strikingly similar. This suggests that the clusters I-L38A and I-L38B probably travelled together to the west. This means that I-L38 carriers were not travelling alone (eg as travelling smiths or metal merchants) but as a group.

On their way to the British Isles the mixed I-L38 lot seems to have crossed Belgium what ex-plains its presence around the rivers Meuse and Scheldt. On the British Isles the trendlines have a weak relation, possibly indicating several (non related) crossings of the Channel.

- The trendlines of I-L38A and I-L38B to

the east and northeast are less similar indicat-

ing an organic gradual diffusion. Seen the distri-bution of the I-L38 subclades around known EBA locations, it is plausible to attribute the spread east to EBA-groups.

This also might explain the predicted presence

of the 3 I-L38 clusters in the Lichtenstein cave (the yellow trendline actually crosses the Lichten-stein location).

Cluster I-L38B seems to have gone further to the east than I-L38A. Conclusions (figure 3):

Since the trendlines only are very rough indi-cators, the comparison of the trendlines of sev-eral clades only leads to hypotheses:

- Three trendlines, I-L38 (orange), R-U152

(dark blue) and R-L21 (light blue), cross near contemporary Frankfurt, suggesting a common origin that could be linked to the La Tène culture that started to spread from the middle Rhine re-gion.

- R-U152 also is connected to Switzerland,

this being the reason Steve Jones classifies this group as Alpine Celtic. The trendline crosses the archaeological site of La Tène that gave its name to this culture.

- The trendline of R-S21, classified as River

Celtic by Steve Jones, partly resembles the trendline of I-L38. This clade also was present in the Lichtenstein cave.

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Results of the geo-analysis of I-L38 subclades

Figure 2: the I-L38 A (yellow), I-L38B (red) and I-L38C (blue) trendlines projected on a map.

Results of the geo-analysis of I-L38 and other haplogroups

Figure 3: the I-L38 (orange) , I-M223 (pink) and R1b-trendlines projected on a map. R-L21 (light blue), R-S21 (black), R-U152 (dark blue).

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Method 2: PHYLIP tree of I-L38

In order to get an idea of the timeframe of the I-L38 division, Tim Weakley made the follow-ing PHYLIP tree using:

the infinite allele mutation model;

average mutation rate derived by Doug McDonald from the Sorenson database;

a probability of 95% that the TMRCA is no

longer than indicated;

an average generation interval of 30 years.

Figure 4: PHYLIP tree, kindly made by Tim Weakley. The interpretation of this tree is entirely to the account of Hans De Beule.

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Interpretation:

It is interesting that the oldest branches (GER1 and GER9) which separated 4000 years ago, cover the German EBA area.

During the dark grey coloured period there

apparently was a direct migration to the British Isles.

When looking at I-L38 Upper Rhine samples

(encircled) and the other I-L38 samples it is re-markable that most splits happened around 2200 years ago (the light grey area), during the late iron age.

Method 3: study of the historical-cultural context

Traditionally the iron age is divided in the Early Iron Age (corresponding to the Hallstatt culture) and the Late Iron Age (corresponding to the La Tène culture).

Around 600 BC trade in continental Europe

started to change, shifting wealth and power to the area west of the Alps. Probably this shift was a result of the founding of the Greek colony Massalia (Marseille). Because of this the trade routes that crossed the Alps were abandoned in favor of new routes that followed the course of the Rhône. In the area west of the Alps (the Southwest of Germany, the Rhine area and East-France) a hierarchical society develops, leaving us impressive grave hills and grave gifts.

Medio 450 BC central Italic cultures took over

the leading role from the Greek colonies. The new trade routes directly cross the Alps, through Switzerland to the Middle Rhine area and to middle France. Grave goods show that men were buried with there weapons and suggest small egalitarian communities of warriors. For unknown reasons this is the period of the Celtic migrations, the most spectacular being the besiege of Rome and the plundering of Delphi (Bourgeois, 2003).

Although the spread of I-L38 has nothing to

do with Celtic migrations to the South, there seems to be a relation between migrations that connect the Rhine area to the west. The early La

Tène migrations left traces along the rivers Meuse, Scheldt and other rivers on the British Isles.

Traces along the river Meuse (Maas in Dutch)

In Baarlo, along the Meuse (where one of the

two Dutch I-L38) samples is located two bronze buckets were found that served as urns.

These buckets indicate a link to the Upper

Rhine region.

Figure 5: distribution of bronze buckets in Europe – Bloe-mers 1991, after Kimmig 1983 -a: 7th-6th century BC,

b: 5th century BC, c: centres of production. Wallony as well as in Flanders, luxurious La

Tène graves have been found together with bronze buckets that were used as urns, for ex-ample in Eigenbilzen near Maastricht. (Clerinckx, 2005).

In Limburg (border area between Belgium and

the Netherlands) graves with valuable grave gifts have been found. The objects are dated 450 BC

Scheldt

Meuse

UpperRhine

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and are related to the Marne-Middle Rhine tradition. It seems that these graves were an expression of (late) Celtic aristocracy (Bourgeois, 2003).

Traces along the river Scheldt (Schelde in Dutch)

In the early La Tène period lots of depositions (swords and other valuables) were thrown in the river Scheldt. These depositions took place less than 10 km from the two located Belgian I-L38 samples (in Zele and Sint-Amands). In Zele also a grave hill was found. Research on pollen showed that the grounds have been grazed; indi-cating cattle and permanent settlement.

Also in the adjacent Berlare La Tène pottery

was found that could be dated precisely to 450 BC (Bourgeois, 2003).

Traces along British rivers Around 400 BC the La Tène culture extents to

Great Brittain and to Transdanubia (the eastern part of Austria and Hungary) (Haywood, 2001).

A remarkable number of La Tène weapons

have been recovered from lakes, rivers and bogs. Some of the finest examples were dredged from the river Thames in and around London. Other rivers have yielded treasures such as the mag-nificent Witham Shield, from the river Witham, near Lincoln, swords and scabbards from the river Nene near Peterborough, and a unique bronze shield found in a former watercourse at Chertsey, Surrey in 1985. Close contact with the continent only appears at the mid fifth century BC; thereafter the British and French traditions diverged, hinting at the beginning of some de-gree of cultural isolation.

The 20 or more daggers of the La Tène type I

(450-300 BC) found in Britain, mainly in the Thames area, can be arranged in a typological sequence lasting until the 4th century BC.

At least 17 swords of the La Tène I type have

been identified from the rivers Thames and Witham, but only two or three have any claim to

being actual imports with Swiss characteristics. What is perhaps most impressive about the Brit-ish swords is their relative isolation from conti-nental development. If the number of La Tène brooches can be taken to be a reflection of inten-sity of importation, then the period from 450 to 350 BC was a time of much interaction, after which, until about 100BC, the intensity of contact dramatically declined (Cunliffe, 2005).

Archaeological evidence of the so called La

Tène Arras group (named after the Arras ceme-tery in Yorkshire) indicates a folk movement into eastern Yorkshire early in the 4th century BC. The evidence suggests small bands arriving with little more than their personal equipment and settling down among the (Bell Beaker) natives. Artefacts show cultural influences affinity to the Alsace re-gion, Switzerland, Champagne and Burgundy re-gion (Cunliffe, 2005).

As the map below illustrates it is safe to say

that the British Isles received a La Tène influx from the Atlantic coast ranging from the estuary of the Rhine/Meuse/Scheldt to the estuary of the Seine and even more south to the estuary of the Loire.

Figure 6: the routes by which concepts of La Tène art reached Britain and Ireland (Kearney, 2006).

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To link the map above to the actual Y-DNA distribution one should take into account that:

- also in later centuries/millennia there was

an influx of continental Y-DNA from the same At-lantic shores;

- the invasions of later centuries (Romans, Angles, Saxons, Jutes, etc) pushed the older «na-tive» populations to the west and to the north. Conclusion

The definition of the La Tène culture as for-mulated in the Columbia Encyclopaedia fits the spread of I-L38 strikingly well:

lä tĕn, ancient Celtic site on Lake Neuchâtel,

Switzerland, that gives its name to the second and final period of the European Iron Age. The earliest phase of Tenian culture, from the 6th to the late 5th cent. b.c., spread from the middle Rhine region East into the Danube valley, South into Switzerland, and West and North into France, the Low Countries, Denmark, and the British Isles; this was the period of the first of the great Celtic migrations. Tenian culture flourished until subjected to the advances of the Roman Empire. Native coinage appeared in Gaul during the latter part of the period, along with the forti-fied townships eventually conquered by Julius Caesar.

I-L38 could have migrated out of the Upper

Rhine area (Rhineland-Palatinate) in the era of La Tène migrations (around 450 BC).

To reach the British Isles I-L38 crossed,

among other regions, the Low Countries. Ar-chaeological artefacts and studies suggest that the migration that crossed the Low Countries took place in three stages:

in a first phase along the Meuse; in a second phase along the Scheldt;

the cross of the Channel in a third phase. Data of the ongoing Hertogdom Brabant DNA

project in Flanders will be welcome to finetune this theory.

The La Tène link does not explain the pres-

ence of I-L38 in the Lichtenstein cave (1000-750 BC) unless there were earlier bronze or iron age migrations to the north-east. Seen the distribu-tion of I-L38 around well known EBA locations it is plausible to presume an early and gradually diffusion of I-L38 from the Rhine into north-eastern direction.

Lingering questions

It still is a mystery why I-L38 is so small in size. The answer to this question might reveal more about the historical position of I-L38 and offers a challenge for future research.

It also is a pity that the exact frequencies of

I-L38 in SW Germany, in Belgium, in Luxemburg, in north-east France, in Austria, Switzerland, north Italy and the British Isles is not available (yet).

Exact frequencies could help solving questions

as: Is there a historical relation between the (low)

frequency in Belgium and the genocides Julius Caesar described in his De Bello Gallico (account of his campaign against the Belgae) -or- Did Roman pressure trigger a migration of Belgae to the British Isles?

How did I-L38 cross France ? What is the relation between English, Welsh,

Scottish and Irish I-L38’s? How and when did I-L38 migrate south to the

Alps (Austria, Switzerland, North-Italy).

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Webreferences 1. PHYLIP Software: http://en.wikipedia.org/wiki/PHYLIP 2. Haplogroup I subclade modals: http://knordtvedt.home.bresnan.net/FounderHaps.xls 3. Haplogroup I predictor: http://members.bex.net/jtcullen515/haplotest.htm 4. About generation interval:

http://www.smgf.org/ychromosome/generation_interval.jspx )

5. To determine decimal geographic coordinates: http://www.begeleidzelfstandigleren.com/aardrijkskunde/losse_animaties/geopositie.html

6. For an overview of the R1b SNP tree: http://www.isogg.org/tree/ISOGG_HapgrpR09.html

7. About «De Zonen van Adam in Nederland»: http://www.barjesteh.nl/DNAproject.htm 8. About: Ken Nordtvedt’s Generation Spreadsheet http://knordtvedt.home.bresnan.net/Generations2.xls

References 1. Bourgeois I., Gelorini V., De Clercq W., Deforce K. & Van

Strydonck M. 2003c: De ijzertijd in Zele (ca. 800 - ca. 50 v.C.): aan de periferie van een veranderende wereld, Tijdschrift van het Verbond voor Oudheidkundig Bode-monderzoek in Oost-Vlaanderen 57, 11-24.

2. Bloemers JHF, van Dorp T (Editors). Pre- & protohistorie van de lage landen. Open Universiteit, De Haan, 1991, 496p.

3. Clerinckx Herman. Kelten en de Lage Landen. Davidsfonds, Leuven, 2005, 293 p.

4. Columbia Encyclopaedia, Sixth Edition, 2004, Columbia University Press.

5. Cunliffe Barry, Iron Age communities in Britain (an account of England, Scotland and Wales from the Seventh century BC until the Roman Conquest, Routledge, Oxon and New York, 2005.

6. De Beule Hans. Origin, Distribution and Migrations of I2b*-Subclades. 18th of September 2008. http://www.familytreedna.com/public/I2b2/default.aspx

7. De Beule Hans. Origins of Hg I-L38 (I2b2) Subclades. 5th of April 2009. http://www.familytreedna.com/public/I2b2/default.aspx

8. Schilz Felix (2006) Molekulargenetische Verwandtschafts-analysen am prähistorischen Skelettkollektiv der Lich-tensteinhöhle. Dissertation, Göttingen.

9. Haywood, John. The historical Atlas of the Celtic World, 2001 Thames & Hudson Ltd, London.

10. Heem - en Oudheidkundige Kring Berlare, 1999 nr. 2 en 2001 nr. 1.

11. Kearney, Hugh. The British Isles, a history of four nations, second edition, 2006, Cambridge, Cambridgde University Press, p. 28.

12. Libber Birgit. Zu den frühbronzezeitlichen Gruppen in Süddeutschland. Universität, Leipzig, 2004.

13. Oppenheimer Stephen (2006) Myths of British Ancestry. Prospect Magazine. Issue 127, October 2006.

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Appendix A: STR values of the Lichtenstein bones Of the Lichtenstein bones the following STR-values are made public:

T\DYS 391 389i 439 389ii 438 437 19 392 393 390 385a 385b ∑ HG

Y1 11 12 11 28 10 15 16 11 13 25 13 17 6 I2b (100%)

Y2 11 12 11 27 10 15 15 11 13 25 13 17 3 I2b (100%)

Y4 11* 12 11* 10 17* 1 I2b (100%)

Y6 11 12 11 28 10 15 16 11 13 24 13 17 3 I2b (100%)

Y3 11 13 12 29 12 15 14 13 13 23 11 14 1 R1b (100%)

Y5 11 13 11 30 11 14 15 11 13 25 11 13 2 R1a (100%)

*: Uncertain

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Appendix B: Decimal geographical co-ordinates

Oldest Known Place of Origin of the Y-DNA

sample

Longitude (pos=east neg=west)

Latitude (north)

I-M38 I-M38A 1695, Martock SOM UK -2,7658 50,9705 1801 Ballycloghan, Antr, N. Ireland

-7,7045 53,6521

1690, Upton, Nottinghamshire

-2,8861 53,2143

1649-1729 Bogen, Bavaria 12,6893 48,9109 °1625 Poppenhausen 10,1467 50,0921 °1720 Heidelberg, Pfalz 8,6785 49,4007 1791 - 1798 Kreis, Kolmar, Posen

9,4955 53,7322

1620-1700 Geisleden 10,1941 51,3532 1861 Bosseborn (Kreis Hoxter)

9,3073 51,7484

Rees Germany 6,3955 51,7619 1560 Zele 4,04 51,0682 Hoogvliet Rotterdam, 1504 4,3586 51,8633 Baarlo, Netherlands 6,0971 51,3307 Kirrberg, Elzas, France 7,0646 48,8219 Hauteville-la-Guichard, France

-1,3021 49,1255

Bellemagny, Alsace 7,0656 47,6773 °1830, Llanelli, Wales, England

-4,1629 51,6791

1819-1905, Colerne, Wiltshire, England

-2,2593 51,4388

I-L38B 1623-1702 Bedfordshire, England

-0,4812 52,1049

1779-1838 Aldby, England 1,2696 52,8524 1854-1920 Haworth, Yorkshire, England

-1,9535 53,8294

1692-1772, Ballinvoher, French Park,IRE

-8,6668 52,6376

°1635 -1704 Wiltshire England

-1,9916 51,246

1490 Wiltshire, England, UK

-1,9916 51,246

1820, MountBellew, Irleland

-8,5004 53,4609

1765, Huntly, Aberdeenshire

-2,4042 51,8718

1715 Methingen Metzingen 7,779 52,3148 °1866 Darmstadt 8,651 49,8718 1832 Berlin 13,4119 52,5222 Kleve, Germany 6,1313 51,7843

St Amands, Belgium 4,2045 51,054 Bâtie Montgascon(38),France 5,5304 45,5801 Fjellestad,b.1812 Norway. 6,7795 58,0863 Warta,Lodz,Poland 16,669 50,454 °1702 Schauffausen SWIT 8,6355 47,6969 Jastrabie, Slovakia 17,1651 48,1395 Primo Gandola, b. ca. 1812, Bellagio, Italy 9,2629 45,9872 1742, Hojrup, Tonder 9,6027 55,5555 Switzerland Kienberg 7,9677 47,4382 1732 in Palatinate, Ger-many, d. 23 November 1792, Paradise, York Co., Pa. 7,7669 49,433 I-L38C °1755 Ommeray Lorraine 6,6937 48,7113 1880, Lumphanan, Aberdeensh Hall, 1813-186 Hanley cas-tle, Worcestershire, England -2,2334 52,0772 °1840-1892 Whitnash, Warwickshire, England -1,5216 52,2703 R-U152 Ryton Gateshead UK -1,7578 54,946 Galway -9,0516 53,2737 Wortegem 3,5109 50,8515 Naila, Germany 11,7085 50,329 Rokytnice Orlickyck Horach 16,4655 50,1646 Baden 9,0852 53 Jaszarokszallas 19,9787 47,6442 Bentz-Stüdlin 8,503 49,4786 Gondiswil 7,8723 47,1456 Vaud 6,5373 46,5609 Salins 5,8806 46,9426 Aunis 0,8327 46,083 Altavilla Monferrata 8,3779 44,994 Santander 3,8067 43,4607 Blanes 2,7925 41,6739 Bad Königshofen im Gabfeld 10,4673 50,2985 Holbeach Lincs England -0,0139 52,8039 Hartest, Suffolk -0,6799 52,1395 Croxton near Thetford -0,7534 52,4226 Esneux 5,5686 50,5337 Zug Suisse 8,5163 47,166 Civenna Italy 9,2722 45,9427 Stange Hedmark Norway 11,1918 60,7174

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Assentoft 10,149 56,4404 Lunde A Denmark 13,153 55,392 Viereck 14,0412 53,5491 Wauchope -3,1302 55,9335 Isle of man -4,5479 54,2358 Stafford -2,1161 52,8049 Meulebeke 3,2884 50,9497 St Germain en Laye 2,0936 48,8986 Triefenstein 9,6029 49,8095 Kusel, Palatinate Germany 12,0728 52,228 Gerlfongen 9,064 48,799 Nancy 6,1826 48,6906 Brülten, Fischental 8,6739 47,4716 R-L21 Esens Ostfriesland Ger 7,6118 53,6478 Poznan (Pol) 16,9254 52,406 Arendal (Nor) 8,7669 58,4593 Vimmerby (Sweden) 15,855 57,6656 Hardaland (Nor) 6,2876 60,2469 Jämtland 'Sw) 14,9593 63,171 Leidschendam-Voorburg (Ned) 4,3585 52,0754 Mulheim 6,8865 51,4268 Bundesbach 7,3774 49,8419 Wurzburg 9,9278 49,794 Neustadt 11,835 49,8246 Weil-der-Stadt 8,8692 48,7517 Baden-Württemberg 9,3502 48,6614 Le bourgneuf-la-fôret -0,9711 48,1635 Brittany (france) 2,9325 48,2019 Drain (Fra) -1,206 47,339 Montussaint (Fra) 6,2924 47,4329 Zurich 8,538 47,3686 Ranville-Breuillaud -0,1165 45,9021 Ranville-Breuillaud, France -0,1164 45,9017 Stuttgart, Germany 9,1807 48,7763 Bundenbach, Germany 7,3774 49,8415 Zurich, Switzerland 8,5383 47,3685 Heihiller -2,1067 49,1916 Gloucester, England -2,2483 51,8662 Sturton-Le-Steeple, Notts. UK -0,8194 53,3457 Britford, Wiltshire, England 1,7518 53,7937 Chalmers, McDonough,IL;Wales -3,5398 52,4486 Inverness, Scotland -4,2313 57,4763 Isle of Bute, Scotland -5,0561 55,8354

Ayrshire/Renfrewshire -4,5427 55,8296 Sligo, Ireland -8,4709 54,2702 Dublin Ireland -6,2672 53,3436 Belfast, N. Irela -5,9299 54,5968 Limerick, Ireland 1865 -8,6266 52,6634 Antrim, Ireland -6,2169 54,7129 County Cavan, Ireland -7,3357 53,948 Littleton, Tipperary -7,738 52,6367 Marans, France -0,9914 46,3082 Ballymoney,Antrim,No.Ire. -6,2781 54,8626 I-223 Århus Amt, DK 10,2124 56,1577 Londonderry, Ireland -7,325 54,9938 Rennertehausen, Hesse 8,6905 51,0261 Erristø, Vejle, Denmark 9,7036 55,5491 Bozec, Czech Republic 17,5954 50,3186 Devon, England -3,2238 50,703 Tanum, Bohuslan, Sweden 11,3384 58,7189 Oberlustadt, Bayern,Germany 8,2635 49,2438 Reedham, Norfolk, England 1,5688 52,56 Llanfair Dyffrin Clwyd Den. Wales -3,4112 51,4197 Melton, Suffolk, England 1,3332 52,1063 Pentrich, Derbyshire -1,4189 53,0684 Swaffham, Norfolk, Eng. 0,6882 52,6465 Osterwald, Germany 7,0341 52,534 Nova Bela,Austro-Hungary 18,1229 47,7439 Dorchester, ENG -2,4331 50,7102 Ilvese, GER 9,6804 54,403 Suffolk, England 0,9714 52,1868 Farlam, Cumberland -2,6943 54,9217 Genarp, Sweden 13,4011 55,5992 Leksvik, Norway 10,6272 63,6722 Inveraray scotland -5,0735 56,2302 Tyrone, Ireland -8,8881 53,2068 Eichwalde, Germany 13,619 52,3729 Buscot, Berkshire -1,6666 51,6753 Essex Worsham 0,668 51,7657 R-S21 Villach, Karnten, Austria 13,849 46,6155 Basel, Switzerland 7,5812 47,5591 Emmental Distr. Switzerland 7,7505 46,9163 Vilkaviskis, Lithuania 23,0363 54,6476 Rypin, Poland 19,41 53,0654 Montauerweide Prussia 4,4861 51,7888

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Vigny, Pontoise, France 1,9275 49,0768 Eisenberg, Germany 11,9011 50,9677 Meppel, Netherlands 6,1957 52,7003 Oudorp, Netherlands 4,7714 52,6323 Seboncourt, Aisne, France 3,4771 49,9534 Gand, Belgium 3,7211 51,0531 Bæk, Denmark 9,6168 55,2904 Smaland, Sweden 14,3332 58,0402 Altona, Hamburg, Germany 9,9637 53,5429 Szentpéterfa, Hungary 16,48 47,0944 Braunton, Devon, UK -4,1611 51,1128 Dublin, Ireland -6,267 53,3438 Lanarkshire, Scotland -3,7033 55,524 Glatonbury Somersetshire England -2,7176 51,1458 Bitton, Gloucestershire UK -2,4592 51,4242 Isle of Pabbay, Scotland -1,4892 50,7286

London, U.K. -0,1259 51,4997 Manchester, England -2,234 53,4801 Upholland, Lancashire -2,7101 53,5152 Argyll, Scotland -5,2379 56,4289 Levens, UK -2,7873 54,2677 East Brent, SOM, England -2,9388 51,263 Dublin -6,267 53,3438 Killingworth Middlesex CT -1,5675 55,0342 Northumberland, England -2,2562 55,3373 Motherwell, Scotland -3,9948 55,7887 Bideford, Devon -4,2069 51,0193 Baltinglass, Ire. -6,7098 52,9414 Duns, Berwickshire, Scotland -2,3422 55,7778 Cumbria, England -2,7971 54,5768 Stepney, Middlesex -0,0426 51,5173 Kelling, Norfolk, England 1,1137 52,9408

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Phylogenetic Relations and Geographic Distribution of I-L38 (aka I2b2) (29th of June 2010 – http://sites.google.com/site/haplogroupil38/)

Hans De Beule

Abstract

The first section of this paper presents a 49 marker network analysis of 64 I-L38 haplotypes. This network is

used to visualize the phylogenetic relations between the 64 haplotypes. The second section maps the geographic origin of I-L38 samples from several public databases.

Whenever possible the phylogenetic relation between samples with known geographic origin was visualized on the map. Calculating the MRCA between these samples creates a hypothetical timeframe to explain the relations. The third section describes the construction of a distribution map of I-L38. The general conclusion is that, at this moment, most evidence points to a relation between I-L38 and the migrations of Late Bronze Age (Urnfield Cul-ture) and Iron Age (Hallstatt, La Tène) people.

1. On the Structure of I-L38 Introduction

Haplogroup I-L38 is defined by the SNPs L38/S154, L39/S155, L40/S156 and L65/S159. In the ISOGG tree its current name is haplogroup I2b2. It is believed to be an ancient clade. Ken Nordtvedt estimated the Most Recent Common Ancestor (MRCA) of all living I-L38s at 4100 years ago.

Until now, no SNP was found to separate

I-L38. Several researchers structured I-L38, into different clusters, using different markers. Ini-tially Ken Nordtvedt separated I-L38 into 3 clus-ters using DYS448:

I-L38A with DYS448=19; I-L38B with DYS448=21;

I-L38C (later called I-L38Scot) with

DYS448=20.

Determination of subgroups of I-L38 is diffi-

cult because the genetic variety within I-L38 is too small to be significant. To gain in-sight in the structure of I2b2 without narrowing down the structure to a limited set of markers, it is useful to calculate a minimum spanning net-work to cluster the samples.

For the first times such a network was made

for I-L38 starting from a 67 marker set.

Method: calculation of the minimum spanning network and clustering of the samples

According to Bandelt (1999) the multitude of plausible phylogenetic trees is best expressed in a network that displays alternative potential evo-lutionary paths. A minimum spanning tree for a set of sequence types connects all given types, such that the total length (the sum of distances between linked sequence types) is minimal. The minimum spanning network serves as a good point of departure to reconstruct the most likely _____________________________________________________________

Received: July 28 2010; accepted: July 30 2010; published: August 7 2010 Correspondence: [email protected]

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tree by taking geographical information into ac-count.

The Median Joining Networks in this paper are

created by Fluxus 4.5.1.6 Software. To create a minimum spanning network for

I-L38, STR values of 64 samples with known 67 marker set were used. All samples were selected out of the FTDNA I-L38 project. By ignoring the multicopy markers (in the 4 FTDNA panels) the 67 marker set was reduced to the following 49 STR loci to create a network:

DYS393, 390, 19, 391, 426, 388, 439, 392,

458, 455, 454, 447, 437, 448, 449, 460, H4, 456, 607, 576, 570, 442, 438, 531, 578, 590, 537, 641, 472, 406, 511, 425, 557, 594, 436, 490, 534, 450, 444, 481, 520, 446, 617, 568, 487, 572, 640, 492, 565.

Appendix A refers to the used samples that

can be found at the FTDNA I-L38 project. In the Fluxus software it is optional to correct

the STR-values with a customized weight (the standard weight is 10 and the maximum weight is 100).

Following Qamar (2002) a weighting scheme

with a five-fold range was used in the construc-tion of the networks. The weights assigned were specific for each haplogroup and took into ac-count the Y-STR variation across the haplogroup in the whole population. The following weights were used: variance 0-0.09=weight 90; variance 0.1-0.19=weight 70; variance 0.2-0.49=weight 50; variance 0.5-0.99=weight of 30 and variance 1.00=weight 10.

To calculate the MRCA between related haplo-

types with known origin (see Figure 3 and Ap-pendix C):

generations of 31 years were used (this is

the median paternal generation interval calcu-lated by SMGF);

the number of mutations between two samples, as displayed in the Fluxus network, was counted;

Ken Nordtvedt’s I2b specific mutation

rates were used. The average mutation rate used to calculate the MRCA is based on 48 markers: 0,002311697 (or 1/432 per generation). This equals one mutation every 9 generations (432/48)-or- one mutation every 279 years (9 generations * 31 years).

Example: When the number of mutations be-

tween two samples is «20», this equals 20* 279 years = 5580 years between the two haplotypes -or- 5580/2 = 2790 years between both involved haplotypes and their MRCA.

Results

Figure 1 shows the minimal spanning network of the 64 I-L38 samples. This network displays the relationship between the 49 (weighted) STR-values and shows the most likely evolutionary tree. The «torso» of the network is displayed in bold red lines. It is interesting to note that:

1. the «torso» has three distinct arms – the

samples tied to these arms only partially fit the traditional STR-subgroups listed in Appendix A.

2. with the exception of the I-L38Scot clus-

ter, there does not seem to be a relation between clusters of samples and geographical origin, sug-gesting the respective I-L38 clusters spread to-gether as a mixed lot;

3. the sample (A7) nearest the crossing of

the three arms of the «torso» is geographically linked to Solothurn (Switzerland) in the High Rhine area, emphasizing thus the relation of I-L38 and the Rhine.

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Figure 1: phylogenetic 49 marker network of 64 I-L38 samples . Nodes represent haplotypes and are proportional to the number of haplotypes representing it. The length of the links represents the genetic distance. The colour of the nodes and the codename

of the sample refers to the clusters as defined in the FTDNA I-L38 project: yellow nodes = I-L38-A; red nodes = I-L38-B; dark blue nodes = I-L38-Scot (I-L38-C); green nodes = I-L38-D; white nodes = I-L38-E; light blue nodes = I-L38-14.

2. Mapping the Geographical Spread Introduction of I-L38

Since the discovery of I-L38 in may 2005 -

then called I(X) - an increasing number of sam-ples with known geographic origin became avail-able on public genetic databases.

In 2005 it was puzzling why I-L38 was that

absent in northern (Scandinavian) Europe. After it became known, in May 2006, that the

bones that were found in the Lichtenstein cave (in Osterode-am-Harz) could be attributed to haplogroup I-L38, it was tempting to conclude that the Harz region was the cradle of I-L38.

In 2008, a map with pinpointed I-L38 samples showed that the Upper Rhine region (Rhineland – Palatinate) has a high I-L38 frequency. Also was proved that this region harbours a high I-L38 cluster diversity (demonstrated by various DYS448 and DYS19 combinations), making it a likely point of origin of the I-L38 haplogroup. (De Beule 2008 and 2009).

Since 2009, gradually, an increasing number

of samples with known East and Southern Euro-pean origin pop up, demanding an updated state of affairs.

This sections pinpoints the publicly available

samples of I-L38 on a map (and shows the phy-

Interesting I-L38-A clus-ter sepa-rated by DYS442 = 11

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logenetic relations between them) in order to meet this demand.

Method

A distribution map of I-L38 samples was made using the known geographical origin of 84 I-L38 samples from the following public STR da-tabases:

Ysearch (Search by Haplogroup / I2b2); The FTDNA I-L38 project;

SMGF, status May 2010 (with search val-

ues: DYS393=13, 426=11, 392=11, 459=8,10, 455=10, 454=12, YCAII= 19,19).

Refer to Appendix B for an overview of the

pinpointed I-L38 locations.

Results

Figure 2 shows the distribution of the samples with known geographic origin.

Some of the pinpointed samples were also

used in the network analysis. When these phy-logenetic related samples are connected interest-ing patterns emerge (see Figure 3):

one can almost see how I-L38 migrated

from the Upper Rhine to the coast of Normandy (France) to cross the Channel to enter England and Ireland;

from the Upper Rhine region there are

also connections to the north (Southern tip of Norway), east (Poland) and south (Spain);

the Lichtenstein cave (yellow dot) is situ-

ated right on the «northern route»;

Most MRCAs go back to Iron Age (Hallstatt, La Tène) or Late Bronze Age (Urnfield Culture) age, suggesting that I-L38 might have been one of the haplogroups that spread on the waves of these cultures. In this respect it is noteworthy that the 3000 year old artefacts that are found in the Lichtenstein cave also belonged to the Urnfield Culture (Schilz, 2006).

It is important to recognize that there are still

vast areas of Europe of which very little Y-DNA data are known. To draw conclusions about the presence/absence of I-L38 in Spain, Portugal, Italy, the Balkan countries, France, etc. more I-L38 samples with known origin are needed.

To overcome this drawback, section 3 con-

structs a predicted distribution map of I-L38.

3. Constructing a I-L38 Distribution Map Introduction

Since most researches simply did/do not test on SNPs determining I-L38, no real distribution map of I-L38 is available. A rough and predicted I-L38 distribution map can be made using a workaround.

Chiaroni (2009) summarizes the structure of

haplogroup I as shown in Figure 4. According to the ISOGG 2010 tree, SNP M436

(aka P215 or S31) defines haplogroup I2b. Itself, I-M436 (I2b) consists of the subclades:

I-M223 SNP- (defined by SNP M223, called

I2b1 by ISOGG); I-L38 (defined by SNP L38, called I2b2 by

ISOGG).

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Figure 2: pinpointed I-L38 samples with known geographic origin.

Figure 3: phylogenetic related I-L38 samples with known geographical origin; the yellow dot represents the Lichtenstein cave. The white numbers indicate the time (years ago) to the MRCA of the two samples.

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In other words:

«Haplogroup I-M436» = «Hg I-M223» + «Hg I-L38»

-so- «Haplogroup I-L38» =

2Hg I-M4362 - «Hg I-M223»

Remark: a part of the I-M436 haplotypes does not belong to either I-M223 or I-L38. This group is labelled I-M436* (aka I-P215* or S31*). I-M436* is believed to have spread lightly and uni-formly over Europe, excluding Scandinavia. I-M436* is ignored in the reasoning above since the light and uniform spread will not affect con-clusions about I-L38 too much.

Figure 4: the structure of haplogroup I according to Chiaroni (2009).

Method Subtracting the I-M223 % value of a country

from the overall I-M436 % of that country theo-retically results in an indication of the I-L38 percentage in this country.

Estimated % of M436 per country

Drawing on various sources, Eupedia (see ref-erences) displays the frequencies of haplogroup

I2b (SNP M436) in European countries. It does not display percentages on the distribution of I-M223 or I-L38.

Eupedia states that: The sample size for each country or region

is at least 100. Italy, Germany, England and Ire-land have over 2000 samples each, France and Spain over 1000, Portugal over 900, Belgium over 750, the Netherlands, Finland and Hungary over 650, Greece over 500.

Haplogroup I-L38, (not mentioned by Chiaroni)

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The division of Italy is as follows: North Italy is everything until Liguria and Emilia-Romagna; Central Italy comprises Tuscany, Marche, Umbria, Latium and Abruzzo. South Italy is everything else to the south, except Sardinia and Sicily, which have been made into separate categories due to their specific history and rela-tive geographic isolation.

The division of Germany is as follows:

North Germany includes the Schleswig-Holstein, Lower Saxony (+ Hamburg and Bremen) and Mecklenburg-Western Pomerania. West Germany is the Rhineland, Hessen and Saarland. South Germany is Baden-Württemberg and Bavaria. East Germany is composed of Brandenburg, Ber-lin, Saxony-Anhalt, Saxony and Thuringia.

Estimated % of M223 per country

Kalevi Wiik (2008) visualized the distribution

of I-M223 on a map of Europe (see Figure 5). Also Chiaroni (2009) published a distribution map of M223 (see Figure 4). Both maps were used to estimate the percentage of I-M223 in the coun-tries/regions mentioned in the Eupedia distribu-tion table (see Table 1).

Results Because the percentages of I-M223 were

based on gradient maps and different researches

were combined there was a need to correct the outcome. Since the I-L38 values in Belgium (cfr. Hertogdom Brabant DNA project) and the Nether-lands (cfr. Zonen van Adam in Nederland) are known, a correction factor could be applied to correct the combined Wiik + Charioni I-M223 values.

Figure 5: distribution of I-M223 (formerly known as I1c) according to Kalevi Wiik (2008).

This corrected I-M223 % could be subtracted

from Eupedia’s I2b (I-M436) percentages to es-timate the percentage of I-L38 per coun-try/region. In some cases this resulted in a «negative percentage», meaning «even less than zero %» presence of I-L38.

Table 1: the last column displays the estimated percentage of I-L38 per country.

Region/ Haplogroup

I2b% Eupedia

I-M223 category Wiik

I-M223 category Charioni

Combined Wiik &

Charioni category

Corrected%

I-M223 %I-L38

Belgium 4,50 8,00 4,00 12,00 3,20 1,30 Albania 3,00 2,00 0,00 2,00 0,53 2,47 Austria 2,00 5,00 2,00 7,00 1,87 0,13 Belarus 1,00 1,00 0,00 1,00 0,27 0,73 Bosnia-Herzegovina 0,50 2,00 0,00 2,00 0,53 -0,03 Bulgaria 1,00 6,00 0,00 6,00 1,60 -0,60 Croatia 1,00 2,00 0,00 2,00 0,53 0,47 Czech Republic 4,00 1,00 2,00 3,00 0,80 3,20 Denmark 5,00 10,00 3,00 13,00 3,47 1,53 England 4,50 6,00 2,00 8,00 2,13 2,37

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Region/Haplogroup

I2b% Eupedia

I-M223 category Wiik

I-M223 category Charioni

Combined Wiik &

Charioni category

Corrected% I-M223

%I-L38

France 4,00 4,00 5,00 9,00 2,40 1,60 North Germany 5,00 12,00 3,00 15,00 4,00 1,00 East Germany 3,00 12,00 2,00 14,00 3,73 -0,73 West Germany 7,00 8,00 4,00 12,00 3,20 3,80 South Germany 3,00 6,00 2,00 8,00 2,13 0,87 Greece 1,50 4,00 2,00 6,00 1,60 -0,10 Hungary 2,50 1,00 1,00 2,00 0,53 1,97 Ireland 4,00 4,00 0,00 4,00 1,07 2,93 North Italy 2,50 2,00 1,00 3,00 0,80 1,70 Central Italy 5,00 2,00 2,00 4,00 1,07 3,93 South Italy 2,50 2,00 2,00 4,00 1,07 1,43 Latvia 1,00 1,00 2,00 3,00 0,80 0,20 Lithuania 1,00 1,00 2,00 3,00 0,80 0,20 Macedonia 0,00 2,00 0,00 2,00 0,53 -0,53 Netherlands 6,00 10,00 6,00 16,00 4,27 1,73 Norway 1,00 2,00 3,00 5,00 1,33 -0,33 Poland 1,00 1,00 2,00 3,00 0,80 0,20 Portugal 3,00 2,00 0,00 2,00 0,53 2,47 Romania 2,00 4,00 1,00 5,00 1,33 0,67 Scotland 4,00 4,00 0,00 4,00 1,07 2,93 Serbia 4,00 2,00 0,00 2,00 0,53 3,47 Slovakia 1,00 1,00 2,00 3,00 0,80 0,20 Spain 1,00 3,00 0,00 3,00 0,80 0,20 Sweden 2,00 1,00 10,00 11,00 2,93 -0,93 Switzerland 3,00 5,00 2,00 7,00 1,87 1,13 Ukraine 1,00 5,00 2,00 7,00 1,87 -0,87 Colour key: 3-4 % 2-3 % 1-2% 0-1% 0%

Figure 6 visualizes the estimated percentages

on a map of Europe. If this rough approach re-flects the distribution of I-L38 in Europe suffi-ciently correct, it means we can expect more I-L38 samples to pop up along the Danube (Serbia, Slovakia, ...) and in Southern European countries as Portugal, Spain and Italy. Remark on Italy:

The distribution of the surnames of the known

Italian samples suggest that they «leaked» into Italy from across Alps. For the distribution maps of the Italian surnames: see Appendix D. These maps suggest that I-L38 has a higher presence in Northern Italy than in Central Italy.

Remark on Portugal:

One can find an indirect evidence of the pres-ence of I-L38 in Portugal at the Iberian DNA pro-ject of FTDNA. According to the Cullen Hap-logroup I predictor the families Ochoa, Baptista, Cardoso and Dos Anos turn out to be members of I-L38-A (probability 38-39%) or I-L38-RecLOH (probability 38-39%). Three of the four families are linked to Cape Verde, a former Portuguese colony.

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Figure 6: estimated % of I-L38 per country on a map also displaying known origins of I-L38 samples. 4. Further Discussion

Looking at the predicted distribution, it be-

comes clear that there is still much to discover concerning I-L38.

At this moment the geographic distribution of

I-L38 samples with known geographic origin, en-riched with:

phylogenetic relations between samples

shown by the network analysis; MRCA calculations;

the estimated distribution of I-L38 in re-

gions of which very few Y-DNA data are known;

points in the direction of a relation between I-L38 and the spread of Late Bronze Age (Urn-field Culture) and Iron Age (Hallstatt, La Tène) cultures.

The Urnfield Culture (1200-750 BC) developed into the Hallstatt culture (750-450 BC) that led to the La Tène culture (450–50 BC).

Given the age and the spread of I-L38 it

seems logical that the migration of I-L38 from the Upper Rhine region to all its current locations can only be explained by successive independent collective and individual migrations taking centu-ries.

From an archaeological point of view, it can

be argued that the distribution of I-L38 played a role in Late Bronze Age (Urnfield Culture) and Iron Age (Hallstatt, La Tène) migrations.

The former could explain the Urnfield-

artefacts that are found in the Lichtenstein cave; the latter could be tied to La Tène (De Beule, 2009) artefacts.

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In this respect it is interesting to look at the distribution of Iron Age stamped pottery in west-ern Europe. It is important to stress that Iron Age stamped pottery only occurs in specific Euro-pean regions. This type of pottery is found in the vast area encompassed by the Rhine, Danube, Marne and Rhône basins, in Armorica, in Cornwall and the western part of Britain, in the Golasecca and Este culture regions and the Alps, in the high Hérault in southwestern France, in the central areas of the Iberian Peninsula along the Ebre River, the Tagus and the Guadiana Rivers. Figure 7 clearly shows this European phenomenon and emphasises its continental origin in the seventh and sixth centuries BC, slowly spreading further south. Some examples of fine stamped grey pot-tery, show that they used the same La Tène models that we find in northern France, the Brit-ish Isles or Germany. The forms of the vases may vary from region to region, but the stamped motifs are very similar.

Such decorated vessels can be found from the

beginning of the early Iron Age on, or in what used to be called Hallstatt, but especially by the sixth century BC and since the beginning of the late Iron Age or La Tène (Gamito, 2005).

In southern Europe La Tène artefacts are also

found in the south of France, the north of Italy, the south eastern Alps and in the Lower Danube region.

From a linguistic point of view the Hallstatt

and La Tène culture is tied to the Celtic-speaking peoples that entered the historical records with the Hallstatt culture. By the end of the Hallstatt period, the Celts had moved outward from Cen-tral Europe in almost all directions: first into France, Spain, and Britain, then southward into northern Italy, and then eastward into the Bal-kans and Asia Minor as the Galatians of the Bible (see Figure 8) (Noonan, 2008).

Further research focussing on the presence of I-L38 in Serbia, Spain, Portugal, Italy, the Galatai region in Turkey, etc… is needed to generate ac-curate data for these regions and to confirm (or reject) the supposed link to Hallstatt and/or La Tène cultures.

We can expect more I-L38 samples to pop up

in Southern European countries and regions along the Danube.

Figure 7: distribution of Iron Age, La Tène linked, stamped pottery (Gamito, 2005).

Figure 8: supposed spread of Celtic language (Noonan, 2008).

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References 1. Bandelt Hans-Jürgen, Forster Peter, Röhl Arne.(1999) Me-

dian-Joining Networks for Inferring Intraspecific Phylog-enies. Molecular Biology & Evolution, 16(1): 37-48.

2. Barjesteh van Waalwijk van Doorn-Khosrovani S., van Gestel AWJM, Plooij FX, Uitgeversmaatschappij Barjesteh van Waalwijk van Doorn en Co’sZonen van Adam in Ned-erland; Rotterdam en Gronsveld, 2008, 405p.

3. Chiaroni Jacques, Underhill Peter A., Cavalli-Sforza Luca L. Y chromosome diversity, human expansion, drift, and cul-tural evolution 20174–20179 _ PNAS _ December 1, 2009 _ vol. 106 _ no. 48.

4. De Beule Hans. Origin, Distribution and Migrations of I2b*-Subclades, 18 september 2008, posted on http://sites.google.com/site/haplogroupil38/

5. De Beule Hans. Origins of Hg I-L38 (I2b2) Subclades, 5th of april 2009, posted on

http://sites.google.com/site/haplogroupil38/

6. De Beule Hans. Early Bronze Age Origin and Late Iron Age (La Tène) Migrations of I-L38, november 2009, posted on http://sites.google.com/site/haplogroupil38/

7. Gamito Teresa Júdice, The Celts in the Iberian Peninsula, Journal of interdisciplinary Celtic Studies, volume 6, 2005, 571-605.

8. Noonan, Michael. Celtic Crossings Lecture. 2008: https://pantherfile.uwm.edu/noonan/www/Celtic%20lecture.IE.pdf

9. Qamar Raheel, Ayub Qasim, Mohyuddin Aisha, Helgason Agnar, Mazhar Kehkashan, Mansoor Atika, Zerjal Tatiana, Tyler-Smith Chris, Mehdi Qasim. Y-Chromosomal DNA Variation in Pakistan. Am J Hum Genet. 2002 May; 70(5): 1107–1124.

10. Schilz Felix. 2006. Molekulargenetische Verwandtschafts-analysen am prähistorischen Skelettkollektiv der Lich-tensteinhöhle. Dissertation, Göttingen.

11. Wiik Kalevi. 2008. Where did European Men Come From? Journal of Genetic Genealogy, 4:35-85.

Webreferences section 1: On the Structure of I-L38

1. Free network software: http://www.fluxus-engineering.com 2. Haplogroup I subclade modals: http://knordtvedt.home.bresnan.net/FounderHaps.xls 3. Haplogroup I predictor: http://members.bex.net/jtcullen515/haplotest.htm 4. Public STR database:

http://www.familytreedna.com/public/I2b2/default.aspx?section=yresults

5. Calculation of the I-L38 MRCA: http://knordtvedt.home.bresnan.net/MRCA%20Ages.ppt

6. Definition of the Upper Rhine and High Rhine: http://en.wikipedia.org/wiki/File:Rhein-Karte.png

7. Average mutation rates for I2b: Relative-m(i) excel file at http://knordtvedt.home.bresnan.net

8. Median paternal generation interval: http://www.smgf.org/ychromosome/generation_interval.jspx

9. Hertogdom Brabant project: http://www.brabant-dna.org/

Webreferences section 2: Mapping the Geographical Spread of I-L38 1. Public STR database: http://www.ysearch.org/ 2. Public STR data-

base:http://www.familytreedna.com/public/I2b2/default.aspx?section=yresults

3. Public STR database: http://www.smgf.org/ychromosome/search.jspx

Webreferences section 3: Constructing a I-L38 Distribution Map 1. Estimated frequencies of European haplogroups:

http://www.eupedia.com/europe/european_y-dna_haplogroups.shtml

2. Eupedia Sources http://www.eupedia.com/europe/origins_haplogroups_europe.shtml#Sources

3. Sorensen Molecular Genealogy Foundation (aka SMGF): http://www.smgf.org/pages/ydatabase.jspx

4. Public STR database: http://www.familytreedna.com/public/I2b2/default.aspx?section=yresults

5. ISOGG 2010 tree: http://www.isogg.org/tree/ISOGG_HapgrpI.html 6. Iberian DNA project:

http://www.familytreedna.com/public/IberianDNA/default.aspx?section=yresults

Webreferences section: Further discussion

1. Noonan, Michael. Celtic Crossings Lecture. 2008: https://pantherfile.uwm.edu/noonan/www/Celtic%20lecture.IE.pdf

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Appendix A: Samples to Create the I-L38 Network (Figure 1)

Most Distant Ancestor Network code

I-L38-A

Patrick Connolly b.c 1808 Bulgaden Co Limerick IRE A1

Owen Ragon, °1797 A2

Andrew Wolfe A3 Peter Lawrence 1774 (NC) -1856 (IN) A4 A5 Christian Deterding, °1792 and 1799 A6

Fridolin Hurbi, °1767 A7 Robert Thadeus McClellan, °1895 TN - A8 Johan Fuchs (Fox), °1784, Prussia, Germany A9 Georg Simon Wehr, °1720, Heidelberg, Germany A10 Evans A11 Samuel Robison about °1765 - 1826 A12

William Barker A13 Hauteville-la-Guichard, France A14

Hugh Bullock A15 Elijah (Wm E.) Butler, °1819, Colerne, Wiltshire A16 Matthew Weakley, °1695, Martock SOM UK A17 James Smith, °1792 - 1868 A18 William Robinson, °1614-1668 A19 Jean Guittard, °1614, Bellemagny, Alsace, France A20

Joseph Price, °1796 A21 Horatio Huggins, Gingerland, Nevis, WI A22

Loran White, °1952 A23 Brooks A24 Samuel Dale, °1801, Ballycloghan, Antr, N. Ireland A25

Thomas Cullen, °1690, Upton, Nottinghamshire A26 John Fortner, °1775, North Carolina, USA A27

I-L38-B

Patrick Bellew, °1820, MountBellew, Irleland B1

John Garrison, °1799 B2 Claude Reynaud, °1723, Bâtie Montgascon, France B3 John White, 1862-1907 Phila PA B4 Enoch Cornelius Seaver, °1834, North Carolina B5 Ommund Ommundson Fjellestad, °1812, Norway B6 Jesse Campbell, °1820, South Carolina, USA B7

Roger Chievre 960 - 1000 B8 Thomas Sivers 1680 -1714 B9 Stone B10 Haworth B11 Foster B12

Elisha Foster, °1766 -1833, b: VT/MA - USA B13 Henry Hutchison B14 Michael Weathers, °1733, Surry Co. VA B15 Francisco Fox B16 Henry Hainer, Ulster Co. NY B17

I-L38-C or I-L38-Scot

McCratic Sc1 John MacKenzie, °17xx M:Dicie Sc2 Williamson Sc3 Wilson McKenny, °1758, of VA Sc4 Samuel McKinney, ° 1840, TN Sc5 Nicholas McKinney, 1822, Franklin Co., AL Sc6 Johnson McKinney, °1764, VA of AL

Sc7

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I-L38-D

Reece Vandever Morrel, °1795, Camden Dist SC D1 Richard Wootten of Warwick, °1614, England D2 Thomas Boucher, °1780 western VA D3

I-L38-E

William Chaffee Shannon, °1876 NY E1 Edward Richardson, °1701 E2 Adalbertus/ Wojciech Tatucha, °1750, Warta, Lodz, Poland E3

I-L38-14

141 Hezekiah Haney, °1770, Halifax Co. NC 142 James Rawls, °1734, Nansemond Co., Virginia 143 Suárez 144 Johannes Böhly, °1702, Switzerland 145 Grantner 146 Joseph Seiler, °1687, Germany 147

Appendix B: Overview of the Pinpointed I-L38 Locations (Figure 2)

Database Family Name Origin

SMGF Standage °1758, Berry, Sussex, England SMGF Gandola °1864, Porleza, Como, Italy SMGF Jackson °1782, Haydock, Lancashire, England SMGF Hansen °1833, Frederikshavn, Hjorring, Denmark SMGF Bennett °1772, Nutwood, Sussex, England SMGF Mezic °1878, Mali Podlog, Slovenia, Austria-Hungary SMGF Spagnotto °1885, Vallo, Italy SMGF Fosson °1855, Magnechilas Ayas, Aosta, Italy SMGF Tavernier °1847, Lille, Nord, France SMGF Fridal °1795, Lundy, Fyn, Denmark SMGF Adam °1879, Leith, Scotland SMGF Garscadden °1879, Glasgow, Lanarkshire, Scotland SMGF Worthington °1812, Ropley, Hampshire, England SMGF Tietjen °1654, Dellien, Bleckede, Niedersachsen SMGF Weinheimer °1837, Antonin, Tarnopol, Galizien Ukraine SMGF Van Hoesen °1582, Huizen, North Holland, Netherlands SMGF Hill °1674, Old Swinford, Worchestershire, England SMGF Dale °1801, Maghadone, Derry, Ireland SMGF Kronenberger °1811, Plock Poland SMGF Bottemiller °1795, Brockhagen, Steinhagen Westphalia in Kreis

Gütersloh SMGF Craig °1813, Prestonpans, Scotland SMGF Heidenreich °1710, Hullhorst, Lubbecke, Westphalia SMGF Grund °1857, Valašské Meziříčí, Moravia, Austria-Hungary SMGF Clare °1886, Newton, Prestwick, Lancashire, England

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Database Family Name Origin ZvAiN Blaas Rees Germany ZvAiN De Booy De Lier Netherlands ZvAiN Meert St Amands, Belgium ZvAiN Spée Baarlo, Netherlands FTDNA Strohmeier °1649 -1729, Bogen, Bavaria FTDNA Saylor / Seiler °1715, Methingen Metzingen FTDNA Ochs °1625, Poppenhausen FTDNA Wehr °1720, Heidelberg, Pfalz FTDNA Krassin °1791 - 1798, Kreis, Kolmar, Posen FTDNA Hartung °1620 - 1700, Geisleden FTDNA Marschall °1755, Ommeray Lorraine FTDNA Zimmer °1866, Darmstadt FTDNA Underwood °1832, Berlin FTDNA Schlenke °1861, Bosseborn (Kreis Hoxter) FTDNA De Beule °1560, Zele SMGF Lems °1504, Hoogvliet Rotterdam FTDNA Brion Kirrberg, Elzas, FRA FTDNA Hauteville-la-Guichard, France FTDNA Guittard Bellemagny, Alsace FTDNA Reynaud Bâtie Montgascon, France

FTDNA Ommund Ommundson °1812, Fjellestad, Norway

FTDNA Tatucha °1750, Warta, Lodz, Poland FTDNA Lehman °1702, Schauffausen SWIT FTDNA Wanchick Jastrabie, Slovakia (Žiar nad Hronom) FTDNA Peder Andersen °1742, Hojrup, Tonder FTDNA Brabazon °1692 - 1772, Ballinvoher, French Park, Ireland FTDNA Mortimer °1635 - 1704, Wiltshire England FTDNA Bassett °1830, Llanelli, Wales, England FTDNA Butler °1819 - 1905, Colerne, Wiltshire, England FTDNA Oldfield °1813 - 186, Hanley Castle, Worcestershire, England FTDNA Evans °1854 - 1920, Haworth, Yorkshire, England FTDNA Furbey °1840 - 1892, Whitnash, Warwickshire, England FTDNA Sawyer 1623 - 1702, Bedfordshire, England FTDNA Hutchinson °1779 - 1838, Aldby, England

(south east of Whitehaven) FTDNA John van Brussel °1849, Veldhoven, Netherlands FTDNA Dirk Gerritz Kors

Dam °1754, Heemskerk, Netherlands

FTDNA Joseph Seiler °1708 in Sembach, Germany FTDNA Patrick Connolly °1808, Bulgaden Co Limerick IRE FTDNA Wendelin Stehle °1717-1785, Bittelbronn, Hohenzollern Zollernalbkreis,

Tubingen, Baden-Wurttemberg, Germany FTDNA Matthew Weakley °1695, Martock SOM UK FTDNA Thomas Rix °1622, Brancaster, England UK FTDNA Thomas Cullen °1690, Upton, Nottinghamshire FTDNA Edmund Rule °1534, Balsham, Cambridgeshire

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Database Family Name Origin FTDNA Patrick Bellew °1820, MountBellew, Ireland FTDNA George

Cruickshank °1765, Huntly, Aberdeenshire

FTDNA William Cornwell °1609, Terling, Essex, England FTDNA Judde °1554, Winterbourne Wiltshire FTDNA Alexander

Farquhar °1880, Lumphanan, Aberdeensh

ysearch Bower Elgin, Moray, Scotland 1750 ysearch Connolly County Limerick, Ireland ysearch Deterding Hannover/Hanover, Germany ysearch Hurbi Kienberg, Solothurn, Switzerland ysearch Kelly Dundalk, Louth, Ireland ysearch More Wick, Caithness, Scotland ysearch Rule Balsham, Cambridge, England ysearch Salvesen Vennesla, Vest Agder County, Norway ysearch Stehle Bittelbronn-Haigerloch, Hohenzollern, Germany ysearch Suarez Garrovillas de Alconetar, Spain

Appendix C: Overview of the Pinpointed Locations with their Genetic Distance and MRCA Calculation (Figure 3)

Location 1 Location 2 Mutations

counted on Fluxus network

Years ago to MRCA

Mountbellew(B1) Bâtie Montgascon (B3) 28 3.911

Warta Lodz (E3) Bâtie Montgascon (B3) 23 3.213

Garovillas de Alconetar (144) Solothurn (A7) 15 2.095 Solothurn (A7) Methingen (147) 18 2.514 Methingen (147) Fjellestad (B6) 14 1.956

Hannover (A6) Bellemagny (A20) 23 3.213

Bellemagny (A20) Ballycloghan (A25) 17 2.375

Martock (A17) Hauteville-la-Guichard (A14) 13 1.816

Hauteville-la-Guichard (A14) Heidelberg (A10) 15 2.095 Heidelberg (A10) Colerne (A16) 16 2.235 Limerick (A1) Upton (A26) 20 2.794 Upton (A26) Warwick (D2) 28 3.911

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Appendix D: Distribution Maps of the known Italian I-L38 Surnames http://www.gens.labo.net/en/cognomi/genera.html

Fosson Gandola Spagnotto

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Haplogroups E1b1b1c1 (M34) and E1b1b1c1a (M84) among Jews. Could Abraham be E1b1b1c1 or E1b1b1c1a?

A.A. Aliev,

D.L. Tartakovsky

Abstract

The present paper clarifies the TMRCA of the Jews of haplogroup E1b1b1c1, the origin of Jews of haplogroup

E1b1b1c1a (M84) and answers the question: «Could Abraham be E1b1b1c1 (M34) or E1b1b1c1a (M84)?».

Retrospect

The problem of the origin of the Jewish carri-ers of E1b1b1 subclades has been paid sufficient attention [1, 2, 3, 4], including our previous pa-pers [5, 6]. Nevertheless, development of DNA genealogy is not standing still, the number of people being tested for Y-DNA are increasing and the amount of new information is constantly growing, which requires a certain correction of previous conclusions.

In brief, a summary of papers [5, 6] reduces

to the fact that different subclades of haplogroup E1b1b1 (M35) have been presented in the Middle East from ancient times. From these subclades, the maximum time of the most recent common ancestor (TMRCA) among Jews has E1b1b1c1 (M34) subclade: 3375±430 years ago. It comes at a time of settling Jews in Canaan. E1b1b1a1, E1b1b1a2 and E1b1b1a3 subclades were included into the Jewish community in later times. It was suggested that in Pre-Jewish times Canaanite carriers of E1b1b1c1 could be among such histor-ically evidenced people as Amorites, Hittites, Phi-listines and Horites, and one of their representa-

tives was converted to Judaism and became the ancestor of the Jewish line of E1b1b1c1.

The specific structure of Jewish sample of

haplogroup E1b1b1c1 (the vast majority of the haplotypes belong to one cluster) makes the TMRCA very sensitive to including of any new haplotype not belonging to that cluster. In a pre-vious paper [5], due to the lack of 37-marker haplotypes, 25-marker haplotypes (the calcula-tion of which does not give enough accuracy) were used. To improve the accuracy of the calcu-lation, for the present paper only the 37-marker haplotypes, which number significantly increased (N=55), were used. Due to the «instability» of the TMRCA of the Jewish E1b1b1c1, the authors emphasize the importance of the confidence in-terval calculated with 95% probability.

Also, the origin of Jews of haplogroup

E1b1b1c1a (M84), which has the largest number of clusters, has not been adequately studied so far. In addition, the simultaneous presence of E1b1b1c1 (M34) subclades among Jews and Arabs contributed to the assumption of belonging «Y-chromosomal Abraham» (a conditional com-mon ancestor of the Arabs and Jews) to one of these subclades. The question of the lifetime of common ancestors of Jews and Arabs of haplo-

_____________________________________________________________

Received: July 28 2010; accepted: July 30 2010; published: August 7 2010 Correspondence: [email protected]

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groups E1b1b1c1 and E1b1b1c1a has not been studied yet.

The aim of this paper is to clarify the TMRCA

of the Jews of haplogroup E1b1b1c1, the origin of Jews of haplogroup E1b1b1c1a (M84) and answer the question: «Could Abraham be E1b1b1c1 (M34) or E1b1b1c1a (M84)?».

The most recent common ancestors of Jews of E1b1b1c1, E1b1b1c1a

For our research we will use Jewish E1b1b1c1 haplotypes from Haplozone E-M35databases. These are the haplotypes belonging to the E1b1b1c1-D1 cluster [7], and one haplotype from the category E1b1b1c1-Miscellaneous [8]. The calculation according to the algorithm [9] shows that the most recent common ancestor of the sample (N=55, 37 markers, the expected modal haplotype in FTDNA order:

15-25-13-10-18-19-11-12-12-13-12-30-16-9-9-11-12-24-14-20-31-15-15-17-17 – 10-10-19-19-15-13-16-18-33-34-13-10), lived 5650±2820 years ago with the probability of 95%.

To determine the TMRCA of Jews of haplo-

group E1b1b1c1a (M84) we used haplotypes of the Jewish clusters E1b1b1c1a*-A [10], E1b1b1c1a*-B [11], E1b1b1c1a*-C [12] from Haplozone E-M35 database. Our calculation shows that the TMRCA of the sample (N=54, 37 markers, the expected modal haplotype in FTDNA order:

13-24-13-10-17-17-11-12-12-13-11-31-19-9-9-11-11-26-14-20-33-15-15-16-17 – 10-10-19-22-16-13-18-18-31-34-13-10) is 4100±1740 years, with 95% probability.

As one can see, both ages exceed the time of

the Jewish invasion in Canaan. It means that at the time of the invasion and conversion of local population belonging to haplogroups E1b1b1c1 and E1b1b1c1a into Judaism, they were two groups of distant relatives. Rather, each of them were inhabitants of one town conquered the Jews.

Could Abraham be E1b1b1c1 (M34)?

To answer this question let us briefly outline the current situation. According to the Bible and the Muslim tradition, Abraham is the distant an-cestor of Cohens and Seyyids, and more — Jews and Arabs. According to historians, (that general-ly confirm the time mentioned in the Old Testa-ment), in particular, [13, 14, 15], Abraham lived about 4000 years ago.

Currently there is no clear opinion about Ab-

raham’s haplogroup — the known characteristics fit just two haplogroups: J1 and J2. Y-DNA tests of Jews and Arabs are largely related to haplo-groups J1 and J2 and revealed that Jews and Arabs have two common ancestors who lived about 4000 years ago: 4200±500 years for hap-logroup J1 and 4375±530 years for haplogroup J2 [16], in other words, roughly in the period of Abraham, and therefore, in the time of the divi-sion of Arabian and Jewish genealogical lines.

Which of the two ancestors is true Abraham?

J1 or J2? The situation is complicated by the fact that among the Cohens and Seyyids both J1, and J2 are presented. According to the study [17], TMRCA of J1 Seyyids is 1300±260 years ago, which, within the confidence interval, corres-ponds to the lifetime of Imam Ali.

However, if one looks at the haplotypes data-

bases, one can see that the role of «Abraham’s haplogroup» is also eligible for haplogroups E1b1b1c1 and E1b1b1c1a: the carriers of these haplogroups are among both Arabs and Jews, in-cluding as a Seyyids [18] and Cohens [19]. How-ever, judging the draft, the number of E1b1b1c1carriers compared to the one of J1 and J2 is much smaller (1% E1b1b1c1 compared to 37% J1e, 19,6% J2a4h, 12,2% J without down-stream subclades, 14,8% R1b1c1 and 15% of 13 other subclades among Cohens and 3% among Seyyids). It makes an exact calculation of ance-stral ages complicated, but, nevertheless, the origin of Cohens is already has been widely stu-died and reported in the literature [20, 21] and is beyond the scope of the current paper, which studies the origin of haplogroups E1b1b1c1 and E1b1b1c1a among Jews. At the same time, the problem of origin E1b1b1c1 Cohens has not been studied yet and is interesting for genealogists.

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The presence of more than one such haplotype may indicate to their non-random nature. For-mally, it gives reason to consider version about E1b1b1c1 or E1b1b1c1a, studied in this paper, as the haplogroup of Abraham.

At the time of writing the paper (July 2010) in

Sharifs DNA Project and Cohen DNA Project there were 3 haplotypes of E1b1b1c1a Seyyids and 2 haplotypes of E1b1b1c1 Cohens, as well as do-zens of Arabian and Jewish E1b1b1c1a and E1b1b1c1 haplotypes [22]. Haplotypes of E1b1b1c1 Seyyids and E1b1b1c1a Cohens are not available yet (but it is not ruled out that E1b1b1c1 Cohens are non-deep clade tested E1b1b1c1a).

The TMRCAs of Arabian and Jewish E1b1b1c1

and E1b1b1c1a (the expected modal haplotype in FTDNA order:

14-25-13-10-17-18-11-12-12-13-11-31 for E1b1b1c1 and 13-24-13-10 - 16-17-11-12-12-13-11-31-17-9-9-11-11-26-14-20-32-14-15-16-17-10-10-19-22 - 15-13-17-19-31-35-13-10 for E1b1b1c1a) are 8080±3890 and 4080±1440 years ago.

Consequently, the Jewish and Arabian lines of E1b1b1c1 divided one from the other several thousand years before biblical Abraham. On the other hand, the common ancestor of the Jewish and Arabian E1b1b1c1a lived in the same histori-cal era as the biblical Abraham. The result de-monstrates the close affinity of Jewish and Ara-bian E1b1b1c1a.

The TMRCA of E1b1b1c1a Seyyids (the ex-

pected modal haplotype in FTDNA order:

13-24-13-10-17-17-11-12-12-13-11-30-9-9-11-11-26-14-20-32 -14-15-16-17-11-9-9-22-16-13-18-20-32-34-14-10-10-8-15-15-7-10-10-8-10 -10-0-21-23-19-11-12-13-17-7-11-26-20-13-13-12-15-10-12-10-11) is 4080±1560 years ago, that more than thousands years over the era of Ali and excludes their Seyyid origin.

The TMRCA of E1b1b1c1 Cohens (the ex-pected modal haplotype in FTDNA order:

13-24-13-10-17-18-11-12-12-13-11-30-19-9-9-11-11-27-15-20 -32-15-16-16-17-10-10-19-22-17-13-18-18-32-33-13-10-10-8-15-15-7-10-10-8 -10-10-0-21-24-18-11-12-13-17-7-11-25-21-15-13-12-14-10-12-10-11) is 440±410 years ago.

Obviously, for such a small sample it is pre-maturely to draw final conclusions, but, according to the present data, their TMRCA does not con-firm their origin from the Biblical Aaron, and the TMRCA of the Arabian and Jewish E1b1b1c1 much older than Abraham’s lifetime.

Conclusions 1) Subclades E1b1b1c1 and E1b1b1c1a were

included in Jewish community during the con-quest of Canaan. With 95% probability the TMRCA of Jewish E1b1b1c1 and E1b1b1c1a are 5650±2820 and 4100±1740 years.

2) Subclades E1b1b1c1 and E1b1b1c1a both

found among Jews and Arabs, including a small number of Cohens and Seyyids. The calculated TMRCAs show that the most recent E1b1b1c1 an-cestor of Arabs and Jews lived 8080±3890 years ago and the most recent E1b1b1c1a ancestor of Arabs and Jews lived 4080±1440 years ago. The most recent common ancestor of E1b1b1c1a Seyyids lived 4080±1560 years ago, the most recent common ancestor of E1b1b1c1 Cohens lived 440±410 years ago.

3) TMRCA calculations show that, according to

formal characteristics, subclade E1b1b1c1 can not claim to be the Abraham’s haplogroup.

4) The most recent common ancestor of the

Jewish and Arabian E1b1b1c1a lived in the same historical era as the biblical Abraham, but a con-tradiction in the calculated TMRCA of Cohens and Seyyids to historical data exclude haplogroup E1b1b1c1a from the contenders for the role of «Abraham’s haplogroup». Despite this, the re-sults indicate close relationship of Jewish and Arabian E1b1b1c1a lines.

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References 1. Coffman-Levy. A mosaic of people: the Jewish story and a

reassessment of the DNA evidence. Journal of Genetic Genealogy 1: 12-33, 2005.

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3. B. Bonnґe-Tamir et al. Maternal and Paternal Lineages of the Samaritan Isolate: Mutation Rates and Time to Most Recent Common Male Ancestor, 2003.

4. P. Shen et al. Reconstruction of Patrilineages and Matrili-neages of Samaritans and Other Israeli Populations From Y-Chromosome and Mitochondrial DNA Sequence Varia-tion, 2004.

5. Aliev et al. Legacy of Ancient Canaanites in the DNA of modern Jews, Proceedings of the Russian Academy of DNA Genealogy, 2009.

6. Aliev A.A. Origin of «Jewish» clusters of E1b1b1 (M35) haplogroup. RJGG Vol.2, No.1, 2010.

7. Cluster E1b1b1c1-D1 – Jewish cluster 8. E1b1b1c1-Miscellaneous haplotype of Spector 9. Дмитрий Адамов. Расчёт возраста общего предка по

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10. E1b1b1c1a*-A cluster 11. E1b1b1c1a*-B cluster 12. E1b1b1c1a*-C cluster

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chal Narratives: The Quest for the Historical Abraham. Valley Forge, Pa: Trinity Press International, 2002.

15. G. F. Hasel. Chronogenealogies in the Biblical History of Beginnings.

16. Клёсов А.А. Какая гаплогруппа была у Авраама – J1 или J2? Вестник Российской Академии ДНК-генеалогии, Том 3, № 2, 2010, февраль.

17. Клёсов А.А., Лугуев Р.Г. Произошли ли сейиды от ге-неалогической линии Пророка Магомета, а Пророк, как и евреи – от Авраама? Вестник Российской Академии ДНК-генеалогии, Том 2, №7, 2009 г., декабрь.

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22. Haplozone Е-М35

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Contents The origin of haplogroup I1-M253 in Eastern Europe Alexander Shtrunov................................................................................... 1 Arabian clusters of haplogroup E1b1b1c1 (M34) Akper Aliev, Dmitry Tartakovsky................................................................ 12 Origin, Distribution and Migrations of I2b*-Subclades Hans De Beule........................................................................................ 14 Origins of Hg I-L38 (I2b2) Subclades Hans De Beule........................................................................................ 33 Early Bronze Age Origin and Late Iron Age (La Tene) Migrations of I-L38 Hans De Beule........................................................................................ 42 Phylogenetic Relations and Geographic Distribution of I-L38 (aka I2b2) Hans De Beule........................................................................................ 56 Haplogroups E1b1b1c1 (M34) and E1b1b1c1a (M84) among Jews. Could Abraham be E1b1b1c1 or E1b1b1c1a? Akper Aliev, Dmitry Tartakovsky................................................................ 72 About the influence of population size on the accuracy of TMRCA estimation, done by standard methods using STR locus complex Dmitriy Adamov, Sergey Karzhavin (Translation - Vasili Gavrilov)................... 76