linguistic and genetic diversity - ddl · genetic flow for l3e (13% in siwan and 3% in beni...

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2005 - Aussois The Berbers Linguistic and genetic diversity J.-M. DUGOUJON and G. PHILIPPSON UMR 8555 CNRS Toulouse UMR 5596 CNRS Lyon

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Page 1: Linguistic and genetic diversity - DDL · Genetic flow for L3e (13% in Siwan and 3% in Beni Snassen) : migration waves from the Horn of Africa ? L1, marker of West and Central Africa,

2005 - Aussois

The BerbersLinguistic and genetic diversity

J.-M. DUGOUJON and G. PHILIPPSON

UMR 8555 CNRS ToulouseUMR 5596 CNRS Lyon

Page 2: Linguistic and genetic diversity - DDL · Genetic flow for L3e (13% in Siwan and 3% in Beni Snassen) : migration waves from the Horn of Africa ? L1, marker of West and Central Africa,
Page 3: Linguistic and genetic diversity - DDL · Genetic flow for L3e (13% in Siwan and 3% in Beni Snassen) : migration waves from the Horn of Africa ? L1, marker of West and Central Africa,
Page 4: Linguistic and genetic diversity - DDL · Genetic flow for L3e (13% in Siwan and 3% in Beni Snassen) : migration waves from the Horn of Africa ? L1, marker of West and Central Africa,

The Berber ‘world’

Page 5: Linguistic and genetic diversity - DDL · Genetic flow for L3e (13% in Siwan and 3% in Beni Snassen) : migration waves from the Horn of Africa ? L1, marker of West and Central Africa,

2005 2005 –– AussoisAussois

Linguistic approach

The Berber language and its place in Afro-Asiatic

Sub-Classification

Characteristics of Siwi

The case of Tuareg

Berber and Afro-Asiatic

Page 6: Linguistic and genetic diversity - DDL · Genetic flow for L3e (13% in Siwan and 3% in Beni Snassen) : migration waves from the Horn of Africa ? L1, marker of West and Central Africa,

2005 2005 –– AussoisAussois

The genetic markers

Gm and Km immunoglobulin allotypes

Mitochondrial DNA (mtDNA): haplogroups and sub-haplogroups

Y chromosome haplotypes

Page 7: Linguistic and genetic diversity - DDL · Genetic flow for L3e (13% in Siwan and 3% in Beni Snassen) : migration waves from the Horn of Africa ? L1, marker of West and Central Africa,

2005 2005 –– AussoisAussois

Gm and Kmimmunoglobulin allotypes

Page 8: Linguistic and genetic diversity - DDL · Genetic flow for L3e (13% in Siwan and 3% in Beni Snassen) : migration waves from the Horn of Africa ? L1, marker of West and Central Africa,

2005 2005 –– AussoisAussois

The populations

Page 9: Linguistic and genetic diversity - DDL · Genetic flow for L3e (13% in Siwan and 3% in Beni Snassen) : migration waves from the Horn of Africa ? L1, marker of West and Central Africa,

2005 2005 –– AussoisAussois

Multidimensional Scaling

Page 10: Linguistic and genetic diversity - DDL · Genetic flow for L3e (13% in Siwan and 3% in Beni Snassen) : migration waves from the Horn of Africa ? L1, marker of West and Central Africa,

2005 2005 –– AussoisAussois

Principal Component Analysis: 1st axis (85%)

Page 11: Linguistic and genetic diversity - DDL · Genetic flow for L3e (13% in Siwan and 3% in Beni Snassen) : migration waves from the Horn of Africa ? L1, marker of West and Central Africa,

2005 2005 –– AussoisAussois

Gradient of Gm haploptypic frequencies

Gm*3;23;5*

Gm*3;..;5*

Page 12: Linguistic and genetic diversity - DDL · Genetic flow for L3e (13% in Siwan and 3% in Beni Snassen) : migration waves from the Horn of Africa ? L1, marker of West and Central Africa,

2005 2005 –– AussoisAussois

Gradient of Gm haploptypic frequencies

Gm*1,17;..;5*

Gm*1,17;..;5*,28

Gm*1,17;..;10,11,13,15,+28

Gm*1,17;..;5,6,10,11,14,+28

Gm*1,17;..;5,6,24,+28

Page 13: Linguistic and genetic diversity - DDL · Genetic flow for L3e (13% in Siwan and 3% in Beni Snassen) : migration waves from the Horn of Africa ? L1, marker of West and Central Africa,

2005 2005 –– AussoisAussois

Gradient of Gm haploptypic frequencies

Gm*1,17;..;10,11,13,15,16

Gm*1,3;+23;5*

Page 14: Linguistic and genetic diversity - DDL · Genetic flow for L3e (13% in Siwan and 3% in Beni Snassen) : migration waves from the Horn of Africa ? L1, marker of West and Central Africa,

2005 2005 –– AussoisAussois

Gradient of Gm haploptypic frequencies

Gm*1,17;..;10,11,13,15,+28 Gm*1,17;..;5*

Page 15: Linguistic and genetic diversity - DDL · Genetic flow for L3e (13% in Siwan and 3% in Beni Snassen) : migration waves from the Horn of Africa ? L1, marker of West and Central Africa,

2005 2005 –– AussoisAussois

Minimal Spanning tree: genetic distances (12 Gm haplotypes)

Page 16: Linguistic and genetic diversity - DDL · Genetic flow for L3e (13% in Siwan and 3% in Beni Snassen) : migration waves from the Horn of Africa ? L1, marker of West and Central Africa,

2005 2005 –– AussoisAussois

Gm allotypes

Homogeneity of the Northern African Berber (and Arab) populations :

~ 20 % of the sub-Saharan haplotypes in all the populations

80 % of the Gm haplotypes frequency in common with Europeans and West Eurasians

IsseqquamarenTuaregs (Algeria) are different from KelNam Tuaregs (Niger)

Siwan, with more than 50 % of sub-Saharan haplotypes are related to Semitic and Couchitic populations (owingto the caravans, as well as the slave market ?)

Page 17: Linguistic and genetic diversity - DDL · Genetic flow for L3e (13% in Siwan and 3% in Beni Snassen) : migration waves from the Horn of Africa ? L1, marker of West and Central Africa,

2005 2005 –– AussoisAussois

Gm allotypes

Clear differentiation of Northern and Eastern Berbers

If the South-North genetic gradient is marked on bothsides from Sahara, the same is not true for East-Westgradient

Page 18: Linguistic and genetic diversity - DDL · Genetic flow for L3e (13% in Siwan and 3% in Beni Snassen) : migration waves from the Horn of Africa ? L1, marker of West and Central Africa,

2005 2005 –– AussoisAussois

Mitochondrial DNA (mtDNA):

Page 19: Linguistic and genetic diversity - DDL · Genetic flow for L3e (13% in Siwan and 3% in Beni Snassen) : migration waves from the Horn of Africa ? L1, marker of West and Central Africa,

L0 L1b L1eL2a

L2b

L3e

L4g

U6 total

M1

U3

U5b

K

HV1

(preHV)1

H

pre*V1

V

J2

T total X W

L0

L1b

L1eL2a

L2b

L3e

L4g

U6 total

M1

N1bU2bU2eU3U4U5b

K

HV1

(preHV)1

H

pre*V1

V

J2

T total

X

W

L0

L1bL1e

L2aL2b

L3e

L4g

U6 total

M1

N1b

U2b

U2e

U3

U4

U5bK

HV1

(preHV)1

H

pre*V1

V

J2

T total

X W

Page 20: Linguistic and genetic diversity - DDL · Genetic flow for L3e (13% in Siwan and 3% in Beni Snassen) : migration waves from the Horn of Africa ? L1, marker of West and Central Africa,

2005 2005 –– AussoisAussois

Mitochondrial DNA

sub-Saharan North African European

and West Eurasian

Moroccan Berbers

Asni (Rhiraya) 22,6 11,3 66,1

Bouhria (Beni Snassen) 13,9 2,8 83,3

Algerian Berbers

Ghardaia (Mozabite) 14 28,2 57,8

Egyptian Berbers

Siwa 24 0 76

Page 21: Linguistic and genetic diversity - DDL · Genetic flow for L3e (13% in Siwan and 3% in Beni Snassen) : migration waves from the Horn of Africa ? L1, marker of West and Central Africa,

2005 2005 –– AussoisAussois

Distribution of the H1 and H3 sub-haplogroups frequencies

Page 22: Linguistic and genetic diversity - DDL · Genetic flow for L3e (13% in Siwan and 3% in Beni Snassen) : migration waves from the Horn of Africa ? L1, marker of West and Central Africa,

2005 2005 –– AussoisAussois

Mitochondrial DNA

H1 and H3 subhaplogroups (coalescence ages ~ 11,000) are the markers of late-glacial expansions ofhunter-gatherers from the Franco-Cantabrian refuge, after the « Last Glacial Maximum », about 20,000 years ago

H1 displays a high frequency among North Africanpopulations (10 to 20 %), with a maximum in Berberpopulations

Only 1 % frequency in Siwa

Page 23: Linguistic and genetic diversity - DDL · Genetic flow for L3e (13% in Siwan and 3% in Beni Snassen) : migration waves from the Horn of Africa ? L1, marker of West and Central Africa,

2005 2005 –– AussoisAussois

Mitochondrial DNA

L haplogroups:

Genetic flow for L3e (13% in Siwan and 3% in Beni Snassen) : migration waves from the Horn of Africa ?

L1, marker of West and Central Africa, is more frequentin Northern African Berber populations (7-9%) than in Siwa (1%)

L4g, marker of East Africans, is only found in Siwa (4 %)

Page 24: Linguistic and genetic diversity - DDL · Genetic flow for L3e (13% in Siwan and 3% in Beni Snassen) : migration waves from the Horn of Africa ? L1, marker of West and Central Africa,

2005 2005 –– AussoisAussois

Mitochondrial DNA

M1 haplogroup

17% at Siwa and 4% at Bouhria (Beni Snassen) and Asni (Rhiraya)

M1 distribution correlates with the spread ofAfro-Asiatic languages (?)

Page 25: Linguistic and genetic diversity - DDL · Genetic flow for L3e (13% in Siwan and 3% in Beni Snassen) : migration waves from the Horn of Africa ? L1, marker of West and Central Africa,

2005 2005 –– AussoisAussois

Y chromosome haplotypes

Page 26: Linguistic and genetic diversity - DDL · Genetic flow for L3e (13% in Siwan and 3% in Beni Snassen) : migration waves from the Horn of Africa ? L1, marker of West and Central Africa,

2005 2005 –– AussoisAussois

Y chromosome haplotypes

Sample tested: Beni Snassen (67), Rhiraya (54) and Siwi (93)

Markers: > 70 biallelic markers (including some new unpublished)

11 microsatellites

20 distinct binary haplogroups

Page 27: Linguistic and genetic diversity - DDL · Genetic flow for L3e (13% in Siwan and 3% in Beni Snassen) : migration waves from the Horn of Africa ? L1, marker of West and Central Africa,

2005 2005 –– AussoisAussois

Y chromosome haplotypes

Close relationship between the two Moroccan Berberpopulations (78-80% E-M81)

E-M81 was found in only one Siwi (1%)

Very low frequency (2-6%) of haplogroups of Europeandescent (such as R-M269, J-M12 and E-M78 cluster α) in allthe berber populations

Relatively low frequency (2-14%) of haplogroupscommonly found in the Middle East (J-M267 and G-M201)

Page 28: Linguistic and genetic diversity - DDL · Genetic flow for L3e (13% in Siwan and 3% in Beni Snassen) : migration waves from the Horn of Africa ? L1, marker of West and Central Africa,

2005 2005 –– AussoisAussois

Y chromosome haplotypes

Beni Snassen and Rhiraya Berbers from Morocco show

relatively low amount of sub-Saharan Y chromosomes, almost

exclusively E-DYS271 (7%)

Siwa Berbers have a similar frequency (6%) of this

haplogroup, but other sub-Saharan haplogroups (e.g. B-M109

and E-V6) have been observed at high frequencies: about 60% on the whole (these haplogroups are very rare north of the

Sahara)

Page 29: Linguistic and genetic diversity - DDL · Genetic flow for L3e (13% in Siwan and 3% in Beni Snassen) : migration waves from the Horn of Africa ? L1, marker of West and Central Africa,

2005 2005 –– AussoisAussois

Y chromosome haplogroups

0.000.100.200.300.400.500.600.700.800.90

Rhiraya Beni-Snassen

Siwa

Berber populations

Freq

uenc

ies Autochthonous

sub-SaharanMiddle easternEuropeannot assigned

Berbers from the North West and North Eastare genetically quite distinct

Page 30: Linguistic and genetic diversity - DDL · Genetic flow for L3e (13% in Siwan and 3% in Beni Snassen) : migration waves from the Horn of Africa ? L1, marker of West and Central Africa,

2005 2005 –– AussoisAussois

Y chromosome haplotypes

Relative high microsatellite diversity in Siwa Berbers suggests that their presence cannot beascribed to recent bottleneck or recent foundereffect

Sub-Saharan gene flow(s) reflect(s) ancientinteractions, before Sahara became dry ?

Page 31: Linguistic and genetic diversity - DDL · Genetic flow for L3e (13% in Siwan and 3% in Beni Snassen) : migration waves from the Horn of Africa ? L1, marker of West and Central Africa,

2005 2005 –– AussoisAussois

Y chromosome haplotypes

Y haplogroup sharing between Berbers andMiddle East – Eastern Africa is very limited

East-African or Middle Eastern origin of theberber ? Y data doesn’t permit to answer thesequestions

Page 32: Linguistic and genetic diversity - DDL · Genetic flow for L3e (13% in Siwan and 3% in Beni Snassen) : migration waves from the Horn of Africa ? L1, marker of West and Central Africa,

2005 2005 –– AussoisAussois

Pastoralism and lactase genetics

In Europe, the putatively causal allele (-13910T) for lactase tolerance has a frequency of ~ 85 %

In sub-Saharan Africa, this frequency is 0 %

In Berbers from North West : 25 %

The distribution of the other haplotypes P,X and Y shows that migrations from the Sahara were limited

Page 33: Linguistic and genetic diversity - DDL · Genetic flow for L3e (13% in Siwan and 3% in Beni Snassen) : migration waves from the Horn of Africa ? L1, marker of West and Central Africa,

2005 2005 –– AussoisAussois

Pastoralism and lactase genetics

The positive selection pressure on lactase suggests that contemporary Berber populations possess the genetic signature of past migration of pastoralists from the Middle East (Neolithictransition)

Page 34: Linguistic and genetic diversity - DDL · Genetic flow for L3e (13% in Siwan and 3% in Beni Snassen) : migration waves from the Horn of Africa ? L1, marker of West and Central Africa,

2005 2005 –– AussoisAussois

Perspectives

Mali (area of Tombouctou): linguistic and geneticinvestigation in progress on Tuareg

Siwa (hypothesis of a « Zenati » peopling)Investigation in Figuig oasis (Morocco-Algeriaborder)

Other Libyan Desert oasis: Augila ?

Afro-Asiatic and Berber origins: M1 mtDNAhaplotype evolution

Page 35: Linguistic and genetic diversity - DDL · Genetic flow for L3e (13% in Siwan and 3% in Beni Snassen) : migration waves from the Horn of Africa ? L1, marker of West and Central Africa,

2005 2005 –– AussoisAussois

Cartes de similarité génétique

Page 36: Linguistic and genetic diversity - DDL · Genetic flow for L3e (13% in Siwan and 3% in Beni Snassen) : migration waves from the Horn of Africa ? L1, marker of West and Central Africa,

2005 - Aussois

FinancementsFinancements

CNRS: Programme OHLL et OMLL (EUROCORES – ESF)

Conseil Régional Midi-Pyrénées