The male lines of the Maghreb: Phoenicians, Carthage, Muslim conquest and

Berbers

Wim Penninx1

2019-06-22

This document is registered at https://independent.academia.edu/wpenninx

Abstract

In this document i analysed Y-DNA from yfull and ftdna from the Maghreb. I conclude that the
present descendants of E-M81 originate from Phoenicia. The majority of males in the Maghreb has a
Y-DNA that descends from a male line ancestor that lived in Phoenicia about 500 BCE. This founding
father effect is extremely strong and a similar founding mother effect is absent. About 20% has a Y-
DNA that originates in Hejaz (Arabia) and arrived at the Muslim Conquest of the Maghreb. The
Berber population has a lower Y-DNA percentage from Hejaz. I see no indication that the present
population has Y-DNA that descends from the population from the Maghreb from the period before
the arrival of the Phoenicians. The Berber and Tuareg languages descend from the Phoenician
language, and the Tuareg language had the least influence from other languages. A timescale
correction of yfull +10% is more likely than a timescale of yfull without correction or a timescale
correction of yfull +20%.

Introduction

In this analysis I downloaded the data of all individual persons in the https://yfull.com/tree
website [1] on February 15 2019 and concentrated on the people from the countries of
Mauritania, Morocco, Algeria, Libya and Tunisia. This resulted in a set of 80 samples, which is
less than 1 percent of the number of samples in the yfull tree. In this document I used the
time estimates of yfull [2] as a base and successive bootstraps to determine accuracies. The
next step was to see which of the data has characteristics of people whose descendants lived
in the Maghreb. This means that two or more Maghreb samples were neighbours in the
phylogenetic tree. I define two neighbours as two (or more) samples that can have a shared
ancestor, while the other samples in phylogenetic tree do not descent from this shared
ancestor. For this comparison we omit the samples that have an unknown origin (since I
don’t know if they are inside of the Maghreb or outside of the Maghreb). It appeared that
we have ten groups of neighbouring Maghreb samples. Since the majority of the present
population of the Maghreb descends from one branch (E-PF2546, part of E-M81), we split
the analysis of E-M81 in two parts: the analysis of E-PF2546 (1) and the analysis of the non-E-
M81 branch splits of the Maghreb (2).

1
Independent Genetic Genealogy Researcher, Delft, The Netherlands
Analysis of E-PF2546

The largest branch in the Maghreb is well known as the E-M81 group; about 60% of the
population belongs to this branch, see Bekada et al (2013) [3]. It has a clear founding father
effect with founding father characteristics as specified in Sikora (2013) [4] for R1b in Western
Europe. The list of branch splits in E-M81 in yfull with more than three subbranches at the
same SNP level in M81 is reported in table 1. The distribution of the branch splits shows a
major founding father effect at the level of PF2546 and his (grand)child branches A2227,
Z5009, Z5013, A1152. On the later population growth of CTS12227 and PF6794-PF6789 I will
come back in the paragraph “Analysis of CTS12227+PF6789”.

SNP at branch split #subbranches yfull ybp

PF2546 20 2216
PF2546-CTS12227 9 1436
PF2546-A2227 7 2231
PF2546-PF6794-PF6789 9 1320
PF2546-Z5009 16 1959
PF2546-Z5009-Z5013 4 1892
PF2546-Z5009-Z5013-A1152 5 1809
All other SNPs 2-3 50-2805
Table 1

To determine the age of E-PF2546, we have data of 20 branches and many subbranches. To
determine the age of this moment in time I used the 20 ages as determined by yfull, and
used bootstrap routines in R [5,6] to determine the time of PF2546, independent of an
assumed distribution. This resulted in a best value of 2217 years before present with a
standard deviation 113 years. The distribution is shown in figure 1.

The M81 branches before and after E-PF2546

E-PF2546 has 20 subbranches. A few of the branches (e.g. PF6794, CTS12227 and A2227) are
part of the founding father of PF2546 in the Maghreb. Many of the other branches are
narrow and have descendants found in a Spain, Portugal, Italy, France, Britain, Lebanon.

In the table below are the members in the ftdna E-M35 project [7] reported (data collected
on June 10, 2019) for which knowledge on the four subbranch splits (as reported in table 1)
is known (A2227, CTS12227, PF6794 and Z5009) and an originating country is reported. In
this case we separated the M81 marker in three parts of the phylogenetic tree. In this table
we report percentages of a certain area which have a similar history and culture. For each
area the number of tested persons the distribution over the three parts of the tree is
reported. These percentages are insensitive to the differences of number of tested people in
an area or culture2. In table 2 we defined only two areas: the Maghreb and the people
outside the Maghreb. The majority of people outside the Maghreb is north of the
Mediterranean and in large majority from Iberia. The number of people in the other Islamic
countries is very small, and is closer to the distribution of the Maghreb then the distribution
of the people north of the Mediterranean.

SNP at branch split A2227 or Z5009 Rest in #

CTS12227 M81
or PF6794

# in ftdna E-M35 project 90 65 107 262

Magreb (TUN+MAR+DZA+LBY+MRT) 57 17 10 84
Others 33 48 97 178

Magreb (TUN+MAR+DZA+LBY+MRT) 68% 20% 12%

Others 19% 27% 54%
Table 2.

The branches that are in “Rest in M81”, so that describe the period before the founding
father effect in the Maghreb are reported with descendants in Iberia, Southern America,
Western Europe, Great Britain and Ireland, Ashkenazi and Sephardic branches. In the “Rest
in M81” group we find only few people from the Maghreb, other Islam countries, as Saudi
Arabia and a few persons in Italy and Malta.

It indicates that it is most likely that the founding father of PF2546 has his paternal origin in
the Mediterranean Sea, but the location of the origin is unclear from this data. No indication
is present that the Maghreb is preferred above other areas. The population of the “A2227,
CTS12227 or PF6794” grew strongly in the Maghreb. The Z5009 group has partial
descendants in the Maghreb, but also outside the Maghreb.

M81 in the Maghreb

[3] measured the main haplogroups of 1789 persons in the Maghreb. The percentage of E-
M81 ranged between 44-67% in the five different countries (Mauretania, Morocco, Algeria,
Tunisia, Libya). If we take the size of the population into account [8], E-M81 is 56% of the
male population in the Maghreb is E-M81. For one of the five countries one can see that it
does not have a representative sample in [3]. In Libya, which has about 5% of the population
of the Maghreb, only 83 persons were reported in the dataset of [3]. 47 of them were from

2
The numbers and percentages of people tested in different areas or cultures is very different. In a
homogeneous culture (e.g. German, or Ashkenazi Jewish) the distribution of this homogeneous group over
haplogroups can usefully be compared, since a distribution of a cultural group in haplogroups is in general
independent of the number of people tested. In this case a small dependency is present, since some people
might have a preliminary result, and the preliminary result might influence the choice to do a follow-up test. In
this document distribution of people in haplogroups is reported in tables 2 and 5. I see no reason that the
conclusions are influenced by choices of follow-up tests.
two Tuareg villages and had about 50% E-M81 and 50% E-M2, which has its origin south of
the Sahara [9]. The Tuareg are a very small percentage of the population of Libya [10]. Since
endogamy is strong in these villages, we also cannot conclude that the E-M81/E-M2 ratio is
representative for Tuareg; in samples of Pereira et al. (2011) [11] the percentage E-M81 was
larger; E-M2 was present, but much less than 50%.

Reguig et al. (2014) [12] determined the haplogroups of 295 Berber-speaking men in
Morocco with 65-83% M81. It had a decreasing gradient from south to the north, so the
largest percentage was further from the Mediterranean Sea. Solé-Morata et al. (2017) [13]
reported a shared ancestor of 2000-3000ybp of E-M81, which was concluded in citizen
scientist groups several years earlier.

Figure 1. In this figure you can see the distribution of the time to the most recent common
ancestor (tmrca) of E-PF2546. In this diagram you can also see the periods of Punic Wars on
the timescale, which were the most important historic events in the Maghreb in the period
600 BCE-200 CE. In the last Punic War Carthage (which was founded by Phoenicians, [14])
was destroyed by Rome [15]. In the diagram you can also see the maps of the Carthaginian
possessions as sketched in [16] and the artist impression map of E-M81 of [17].

Quality checks: some checks were executed: 1) in case the Spanish Sahara as part of the
Maghreb was added to the sample, one sample was added to the selection, which is below
PF2546, so consistent with the overall result. 2) in case we look at the “research samples” in
yfull, it appears that the distribution of these samples is consistent with that of the overall
analysis. The same is the case for the samples of individual persons that were added after
the moment of download.
Analysis of the non-E-M81 branch splits in the Magheb

In this paragraph I had a look at the nine branch splits in the Maghreb that are outside the E-
M81 branch. One of these nine branch splits had a tmrca was within the 90% error range of
the tmrca of E-PF2546 (see above). It is possible that this branch split (E- FGC18960) was part
of the same founding father effect in the Maghreb. The other eight branches had 8 values of
tmrca outside the 90% error range of the tmrca of E-PF2546, see table 3. It is possible that
the branches are independent splits, but the other option is that they have a shared origin.
The error ranges do not exclude a shared origin. If we assume they have a shared origin, we
can use the same method as I used for E-PF2546 (bootstrap of 8 values). This resulted in a
best value of 1324 years before present with a standard deviation 84 years. The result is
showed in figure 2.

Figure 2: Distribution of bootstrap tests with the time estimates of the branch splits of the
eight Maghreb branches, that are not the result of the founding effect that resulted in the E-
M81 Maghreb branches. In this diagram you can also see the period of Muslim Conquest of
the Maghreb on the timescale [18], which was the most important historic event in the
Maghreb in the period 200 BCE-1000 CE.

The eight branches have the following characteristics.

SNP at branch split Character. Neighbor. yfull yfull

Banches in ancestral (ybp)
yfull (ybp)

V1174 (in E-V65) Mediterr. 2200 1477

FGC7393 (in J1- Y5320) Arab. 1660 1412
ZS4753 (in J1-Y9272) Arab. 1555 1271
ZS5010 (in J1- Y5320) Arab. 1660 1387
 FGC43126 (in J1- Y5320) Arab. 1660 1071
BY130638 (in J2) Jew.+Qatar 11400 1418
M318 (in J2) Jewish Mediterr. 2600 1682
L271 (in J2) Middle East 5700 877
Table 3

Analysis of CTS12227+PF6789

The branches of E-CTS12227 and E-PF2546 are both descending from E-PF2546 and have a
founding father pattern that is not part of the founding father effect of E-PF2546 (see
paragraph “Analysis of E-PF2546”). Since the time estimate of these branches is consistent
with each other they could have the same origin. Similar as for the previous paragraphs we
determined a bootstrap test. The result is shown in figure 3.

Figure 3: Distribution of bootstrap test of the time estimates of 16 branches of E-CTS12227

and E-PF6789. The ancestral neighbours and descendants presently live in majority in the
Maghreb. In this diagram you can also see the period of Muslim Conquest of the Maghreb on
the timescale [18], which was the most important historic event in the Maghreb in the period
200 BCE-1000 CE.

Analysis of all J1 branches in the Maghreb

The second largest haplogroup in the Maghreb is J1; about 15% of the population belongs to
this haplogroup [3]. In the different countries the values range between 0-22% (Libya had
the lowest percentage and Algeria the highest). In the yfull dataset I reported already the
four J1 branches in the paragraph “Analysis of the non-E-M81 branch splits in the Magheb”.
In the yfull dataset we have four J1 samples without a neighbour from the Maghreb. They
have the following characteristics:

SNP at branch split Character. Neighbor. yfull yfull

banches ancestral (ybp)
(ybp)

Y8537 (in J1-Y5320) Arab. 1660 1395

Y12363 (in J1-Y8537, in J1-Y5320) Arab. 1395 1500
S9517 (in J1-Y9272) Arab. 1683 1763
FGC1713 (in J1-Y9271) Arab. 1848 1869
Table 4

The accuracy of the determined ages is limited. The mathematical method results
sometimes in a in some situation where the determined ancestral age is younger than the
determine age of the branch itself. The shared ancestor of all measured J1 Maghreb persons
is Y9271, which has a tmrca of 1848 ybp. It is the largest number of members in the yfull J1
project with a tmrca after 0 CE. We divided the members of J1 in three groups defined on
three SNPs and followed the same method as specified in footnote 2. The first group is the
branch that is defined by the maker Y9271; the second is defined by the marker Y4348,
excluding the members below Y9271; the third is defined by M267 (J1), excluding Y4348.
Y4348 was chosen since it is thought to be the marker that defines the migration from the
Levant to the Arab countries and was the ancestor of the majority of Arab J1 descendants.
The population growth of this branch is large. Characteristics of the three groups are shown
in table 5.

SNP at branch split Y9271 Y4348, M267, #

excl. excl.
Y9271 Y4348

calculated tmrca in yfull (ybp) 1848 3519 18200

# in yfull branch 562 343 370

Arabia (BHR+OMN+ARE+QAT+KWT+SAU) 55% 32% 13% 904

South Arabia +East Africa 28% 41% 30% 46
(SDN+EGY+YEM+TCH+ETH+ERI)
Magreb (TUN+MAR+DZA+LBY) 100% 0 0 25
SYR+JOR+PSE+IRQ+IRN+TUR 34% 37% 29% 79
RUS+AZE+KGZ+ARM+KAZ+GEO 0 3% 97% 98
Ashkenazi (UKR+POL+LTU+SVK+HUN+HRV+CZE+BGR) 0 3% 97% 36
PRT+ESP+COL+MEX+PER+ITA+MLT 0 0 100% 48
AUT+DEU+FRA+GBR 0 0 100% 21
UNK+ALB+GRC 0 0 100% 6
FIN+NOR 0 0 100% 5
IND+PAK 0 0 100% 3
Table 5
Table 5 indicates that the branches Y9271 and Y4348 are Arab branches with hardly any
descendants outside Muslim countries. Branch Y9271 with a tmrca near 150 CE has more
than 50% of the measured J1 Arabs. The differences between the three groups are large and
suggest that the Maghreb descendants descend from ancestors in the heart of the Arab
population, which was likely in the region of the heart of the Hejaz. Notice that the period
between the ancestor of Y9271 is estimated as 150 CE, while Muhammed was born near 570
CE, only about 420 years later. The Umayyad caliphate that took power in Maghreb
originated in the Hejaz.

Maghreb Y-DNA percentages and origin

In this paragraph we use the haplogroup percentages of [3] and corrected them according to
the size of the Maghreb countries to determine an overall percentage for the male lines of
the Maghreb.

SNP at branch split Likely %

origin

A, B, E-M33, E-M2, E-V12, E-V22, R1b-V88 Sahara 12.0

E-M35*, E-V65, E-M81 Phoenician 59.9
J1, J2 Arabs 16.8
E-V13, R1b-M269 Europe 4.8
E-M78*, E-M34, F, G, I, K, P,R R1a, T ? 6.4
E-V32, L, O - 0
Table 6

The percentages of the first category (Sahara) have a large diversity in the different
countries. Percentages above 1 percent are E-M33 (1.5%), E-M2 (5.9%), E-V22 (1.5%), R-V88
(2.3%). The ranges in the different countries are large, resp. 0.0-5.3% (max. in W. Sahara-
Mauretania), 0.7-38.6% (max. in Libya), 0.0-3.0% (max. in Tunisia), 0.9-6.8% (max. in W.
Sahara-Mauretania). The large percentage of 5.9% of E-M2 is partly the result of the sample
selection in [3], see the paragraph “M81 in the Maghreb”. The percentages of European and
Sahara origin are probably the result of slowly neighbouring exchange of small amounts of
people or specific endogamous groups with different history. The Tuareg in Libya are
probably an example of a specific endogamous group (see paragraph “M81 in the
Maghreb”). Since the population in the Maghreb is stratified [36], and the stratifications are
partly related to history, the percentages of the haplogroups are likely different in different
groups in the Maghreb. These values are an overall average in the Maghreb.

Publications on ancient DNA in the line of E-M81

Only few reports of ancient DNA samples in haplogroup E are reported in the literature.
Harney et al (2018) [19] reported one E-Z830 from the Late Chalcolithic, 4500–3900/3800
BCE in Peqi’in Cave, Israel. Lazaridis et al (2016) [20] reported two E-Z830 (M123-) from
Natufian, 12000-9700 BCE in Raquefet Cave, Israel. The other reported samples from the
Levantine are from haplogroup T (nine samples in the Peqi’in Cave, [19]) and haplogroup J in
the Bronze Age ([20] and Haber et al. 2017, [21]). [19] considered that the high percentage of
haplogroup T in the Peqi’in Cave was a result of a strong homogeneity, not of a large
percentage of haplogroup T in the Levant. The branch of E-M81 and E-FGC18960 are in two
descending lines of E-L19, which is parallel to E-Z830 and is one of three main branches of E-
M35 with a tmrca of 24000ybp (the third main branch is E-L539, which is often reported as
E-M78, since it is the most often measured marker). A presence of haplogroup E-L19 in the
region and period of the Phoenicians seems possible.

Ancient DNA from Maghreb was reported in two recent articles. Van de Loosdrecht et al.
(2018) [22] reported DNA from Taforalt, 15000ybp. The Y-DNA analysis was not reported in
details that can be reused3.

Fregel et al. (2018) [23] measured fairly recent DNA from different sites in the Maghreb. Y-
DNA samples were found with haplogroup G and T and two samples with haplogroup E. The
haplogroup E samples were found in Ifri n’Amr or Moussa (near 5000 BCE). The measured
markers are compared with the branches of Poznik et al. (2016) [24]. IAM.4 is in the L19
branch, but has no markers measured in the branches with presently measured living
descendants. It was possible to measure more markers in IAM.5, and all markers of L19 were
measured positive and 16 markers were measured in the M81 branch. 15 were negative and
one was positive. It means that IAM.5 descended from a side branch of the M81 branch that
split of near 13800-(13800-2800)*1/16, which is 13000 ybp (using standard yfull time
estimates).

On the relation between ancient DNA of M81-branch in Ifri n’Amr or Moussa and the
present M81-descendants in the Maghreb

One can consider two scenario’s the ancient IAM.4 and IAM.5 samples of [23]. In the first scenario
the present M81 in the Maghreb is the result of the migration of the M81 branch that split off near
13000 ybp, of which a descendant (IAM.5) was in Ifr n’Amr or Moussa near 7000 ybp (5000 BCE). The
other scenario is that the shared ancestor had several branches in a large geographical area, and the
branch that lived between 13000-7000 ybp was not geographically close to the branch that was the
ancestral line of the present descendants of M81 in the Maghreb (13000-2800 ybp). I favour the
second scenario for different reasons: 1) I do not consider the relation between the M81-branch in
IMA.5 very strong since: 1a) 13000 ybp was before agriculture, organisation and religion resulted
collaboration in larger communities. Human communities were still very mobile. 1b) E-M35 has 12
branches near 13000 ybp with presently living descendants. These branches have a geographical
central locations ranging from E-V13 on the Balkans, E-V22 in southern Egypt and E-M81 in the

3
Six samples were measured. Four have M78 measured according to the table S16, which is indicated as
E1b1b1a1. In the text it claims five were measured, and in the table S16 sample TAF009 is reported as
E1b1b1a1b1, but no measured marker is reported for this branch. The sixth sample has less markers measured.
It is unclear if not reported markers are negative or have insufficient quality and could not be determined. M78
is one of the three main branches of E-M35 (see above), and parallel to the branches of E-L19 and E-Z830.
Maghreb. We find no relation between the geographical locations and their subsequent branch
splits. 2) Strong founding father effects (like E-V13, E-V22, R-P312, E-PF2546) are thought to be the
result of a migrating group arriving with new knowledge in a new area. This pattern is in general
different from in-place population growths, where in-place population growths are smaller in
number of branches, which fits with the pattern that they have to compete with other in-location
families. 3) The patterns of migration and colonies where new descending line arrive at a different
locations were common in this period. Examples of migrations (where contact with original location
is lost) are R-P312 (arriving in Europe) and likely E-V13 (arriving in the Balkan) and R-V88 (arriving at
the south side of the Sahara) typically took place from the Middle Eastern area to other areas in
periods of 8000-5000 ybp. Examples of colonies (where contact with original location is maintained,
and were partly based on trade of products from different regions) are Greek colonies (E-V13 and
probably some J2 branches) and Phoenician, which took place in the period of 3000-2000 ybp. The
present Mediterranean haplogroup distribution is probably strongly determined by these colonies,
where the arriving population was large, knowledgeable and powerful in comparison with the
original population. 4) Given the number of descending branches, it must be related to a powerful
population. The descendants of M81 are distributed on different areas of the Mediterranean in a
short period of time which fits with a thalassocracy, not with a tellurocracy. This means that we have
no reason to expect a strong relation between the location where most of the present descendants
live and the location of origin. 5) As shown in paragraph “The M81 branches before and after E-
PF2546” (table 2) the branches before E-PF2546 have very few people from the Maghreb. It is
unlikely that this small group originated in the Maghreb and had first an expanding population north
of the Mediterranean Sea and later an expanding population in the Maghreb.

The yfull age normalization

The statistical uncertainty is smaller than the potential systematic error. A systematic error
might be the result of the use the time estimate of yfull [2], which uses the estimate of
Poznik et al [25]. The timescale depends on the SNP mutation rate, and different researchers
have used different mutation rates. In general: the mutation rate in Poznik is considered as:
possibly correct, but might by about 20% larger. A first comparison between the time
estimates of yfull and other data is present in the founding father of R1b in Europe at the
arrival massive migration from the steppe and the arrival of Indo-European languages in
Europe, see Haak et al. (2015) [26]. [26] report “suggest that they spread into Europe from
the East after 3,000 BCE”. This is based on C14 measurements of ancestral skeletons. The
founding father branches of R1b (A8039, A8053, P312, S1200, U106) are reported in yfull as:
4400ybp, 4300 ybp, 4500ybp, 5000ybp and 4700ybp respectively. This gives a founding
father of R-L151 of 4800ybp at the start of the increased population growth. P312 has 40
branches, so good statistics. The two independent estimates (yfull of present descendants
and C14 of skeletons) suggest that the systematic error in yfull is less than 10% (500 years).
In table 6 I report the values of the three scenarios: yfull timescale as based on Poznik; yfull
timescale +10% and yfull timescale +20%.
 Origin of the SNP at branch yfull Poznik yfull timescale yfull timescale
Maghreb split +10% +20%
population
growth

Phoenicia PF2546 217 BCE +-113 439 BCE +-125 661 BCE +-136
Arabia non-E-M81 676 CE +-83 543 CE +-90 410 CE +-97
branches
In place growth CTS12227+PF6789 593 CE +-133 452 CE +- 146 311 CE +- 159
Origin of Corded
Ware culture
Yamnaya region P312 2600 BCE+-128 3060BCE+-141 3520BCE+-153
Table 6

Four examples of population growth are the estimated timescales in three different
scenarios: yfull timescale [2] as based on Poznik et al. (2013) [25], and a 10% and 20%
increase of the timescale. The related historic events with relative accurate time estimates
are: 1) Foundation of Carthage (according to Timaeus of Taormina): 815 BCE; 2) Colonies of
the Phoenicians in the 4th century BCE: Lebanon (4 cities), Cyprus (1), Malta (2), Sicily (2),
Sardinia (4), Ibiza (1), Spain (8), Maghreb (20) [16].; Punic wars: 264-241 BCE, 218-201 BCE
and 149-146 BCE; Muslim conquest of the Maghreb: 647-709 CE; Mass grave after the arrival
of the R-P312 in Europe: 2880–2776 BCE [27], which must have occurred after the arrival of
the Corded Ware culture that came from a population that was related to the Yamnaya
culture. The accuracy of this last period (95 percent) is determined by C14 method.

The yfull tmrca’s indicate the birth year of the persons involved. Many of the historic dates
are dates of events, while the people involved have a range of ages. In the case of a
migration of a group of people, one can image that the people in the group are family
related to each other. This would imply that a shared tmrca can be longer ago than the
moment of a migration. At the same time it is also possible that statistical effects of a small
group growing to a large population, will have a tmrca after the moment of migration.

The accuracy of the C14 method and historical dates in the Maghreb are relative accurate.
The statistical variability in these datasets is small if we compare it to the systematic
uncertainty of the yfull historical timescale. In the comparison of the above values, the 0%
correction can be rejected as based on the age of the R-P312-marker. A value of 5-10%
correction can explain several detailed aspects. The period that the people from the Corded
Ware culture conquered Europe was probably short. The population expansion was
extremely strong of R1b at arriving in Europe, and decreased already after the first SNP-
marker, both for R-P312, R-U152, R-DF27, R-DF13 (and I-DF29) given the decrease of
subbranches in the yfull tree. It is unlikely that this took more than the period of two SNP-
markers as maesured by yfull (288 years). This makes a correction of 20% an unlikely
scenario. The branches that are most populous of M81 in the Maghreb (A2227 and Z5009)
show that the population increase is extremely large in this region. Some subbranches of
Z5009 are in the Maghreb, while others have a large percentage in Iberia. This fits easier
with a scenario where the Z5009 marker has an origin before the last Punic war than after
the last Punic war, since the exchange of Phoenician people between Iberia and the
Maghreb is a natural behaviour in an expanding trading nation like Phoenicia, but unnatural
after the conquest of the Romans. A timescale correction of yfull +10% is more likely than a
timescale of yfull without correction or a timescale correction of yfull +20%.

Female lines in Maghreb

The ftdna dataset has reported 178 mtDNA samples of Mauretania, Morocco, Algeria,
Tunisia and Libya (May 8, 2019 [28]). All of these samples have done a complete
measurement of the mtDNA, and are positioned in the phylogenetic tree structure of
https://www.phylotree.org (van Oven and Kayser, 2009 [29]). The largest set of mtDNA of
the Maghreb with the same mtDNA haplogroup is only 5 samples, so the largest group is less
than 4 percent. In most cases the ftdna mtDNA haplogroups have a large mixture of origins;
the Maghreb mtDNA samples are a small percentage of its haplogroup, and the tmrca of
almost all of these branches is much larger than the tmrca of E-PF2546; see Behar et al.
(2013) [30]. The mean time between two mutations on the mtDNA is about
1/(16569*2.5*10^-8) which is about 2400 year. The male Y-DNA shows a strong founding
father effect (about 60% of Maghreb Y-DNA has E-PF2546); a similar female mtDNA founding
mother effect is absent (at most 4% of Maghreb mtDNA has an equal haplogroup with a
tmrca of that of PF2546). The cultural interaction between males and females in the
Maghreb in the last 2500 years resulted in an extreme difference in characteristics in present
variability of male and female haplogroups. In human populations where a similar cultural
interaction is present, male lineages coalesce significantly more recently than female
lineages, see e.g. the discussion in [25].
[3] analysed the mtDNA of a much larger dataset (2426 samples from Maghreb). Since they
did not measure the complete mtDNA, the data cannot be used to determine an upper limit
to a possible founding mother effect. Their data is consistent with the here reported lack of
a strong founding mother effect.

Most likely history (Occam’s razor)

When the Phoenicians migrated over the Mediterranean to the west and established their
the city of Carthage, they had people with Y-DNA E-M81, and some people with E-FGC18960
and E-V65. Philip K. Hitti (professor in Semitic languages) explained that they named
themselves Mauri (“presumably of Phoenician origin meaning western“) in his book “History
of the Arabs” [31]. Lorenzo Nigro showed that all known migrations in the 3rd-1st millennium
BC in the Mediterranean always went from east to west [32]. The colonies were self-
sustaining and descendants live in Iberia and other Phoenician areas. The Phoenicians
created the city of Carthage and a large reign was established. In the area of the Maghreb
only a small amount of people lived, and the advanced knowledge of the Phoenicians and
sufficient quality of agricultural land gave rise to a fast expansion of the new population in
the Maghreb. The expanded population lived both in cities and in the agricultural areas and
as nomads. The cities and agricultural areas were self-sustaining [33]. This resulted in the
founding father effect of a few branches of E-PF2546 and some members of E-FGC18960
(near 500-200 BCE) in the large area of the Maghreb. The language they spoke was
Phoenician which later evolved in dialects and languages that we now know as Berber
languages. The Berber language family and the Semitic language family are thought to be the
closest languages in the Afroasiatic language family [34]. A conclusion of a migration of R1b-
V88 that is likely related to the Afroasiatic language Chadic was drawn earlier [35]. The age
distance between the Chadic languages and the Semitic/Berber languages is consistent with
a departure of the R1b founding father effect of the Chadic R1b-V88 (R-Y7771, 5100 ybp
using standard yfull time estimates [1]). Other members of E-M81 and E-V65 dispersed as
Phoenicians from the Levant (or from Carthage to Mediterranean countries like Italy and
Iberia). Others stayed in the Levant and descendants still live there (Lebanon or Palestine)
and some left the Levant in the Jewish diaspora.

It is likely that the populations of urban areas and Berber areas became separated in the
period 200 BCE-600 CE. Each developed their own style of customs, dialect, and method to
survive, having families and communities [36].

Strabo (near 17 CE, [37]) describes the different nomadic groups in the Sahara; Masaesylians
were the Phoenician nomads in the desert and were very similar in clothing as the Libyans in
the urban areas; they used small but swift horses. The Pharusians and Nigretes, living near
the Aethiopians and probably from black African descent [38] and had bows and scythe-
bearing chariots. Strabo tells us that the groups did not mingle. It is likely that the population
of the Phoenicians expanded and the black Africans did not survive in large percentages in
this period in the Maghreb. In the present Tuareg we see an example of a mix between
people of a majority of E-M81 and a minority of E-M2 African descent. The Tuareg language
is a member part of the Berber language family [39]. Some scholars have claimed before that
the traditional written Tuareg language (Tifinagh-Tamazight) is "one of the oldest scripts in
the world, being close both linguistically and in alphabet to Old Phoenician” [40]. “The
Tuareg languages are generally acknowledged as being the most conservative forms of
Berber speech”. This is consistent with the least influence of other cultures after the arrival
of Phoenician language in Tuareg population [40] . Other Berber and Maghreb populations
received a larger influence of the Arab language.

The Muslim conquest of the Umayyad in the Maghreb and Iberia resulted in an upper class
of Arabs from the region of Hejaz in the Maghreb. The Arabs named the region Maghreb,
meaning “West” in the Semitic Arab language. Again, they brought new knowledge and
again a fast expansion of the population took place in the Maghreb. The arrival resulted in 8
branches of J-Y9271 in the dataset, and four of these J-Y9271 branches show expansion in
this dataset in this period. In the same period the population that had arrived in the
Phoenician period also benefitted of the new knowledge and several of these branches show
expansion (E-V1174 (in E-V65) and E-CTS1117 and E-PF6789 (both descendants of E-
PF2546)). Similar as in the Iberia, Jews arrived with the Arabs and took positions specific
tasks in the society, using their specific knowledge and under the rule of the Arabs. The
branch J-M318 (in J2) that has descendants in Tunis is an example of that. For two branch
splits the branch splits leave more options for interpretation. J-BY130638 (J2) has old
neighbours from Qatar and Jews. Since the ancestral branch is estimated at 11400 ybp,
many options are open. The Tunis branch J-L271 (J2) has a tmrca of 900 ybp, with an
ancestral tmrca of 5700 ybp and old neighbouring branches in the Middle East, but also India
and Sri Lanka. Again, many options are open for this branch. The Umayyad Arabs brought
the Arab language in the Maghreb. It replaced the Phoenician Semitic language in the urban
areas.

The percentage of E-PF2546 remained large in the Berber population, while the percentage
in the urban areas decreased more as a result of the arriving Arabs with haplogroup J-Y9271.
The original Phoenician language remained strongly present in the Berber population of the
Maghreb.

Conclusion

About 60 percent of the male lines in the Maghreb are the result of a founding father effect of
arriving Phoenicians in the Maghreb. About 20 percent of the male lines are the result of arriving
Arabs at the Muslim conquest of the Maghreb. Very small percentages come from Europe and small
percentages have their origin from the African population. It is likely that the European lines arrived
after the arrival of the Phoenicians. No indication of descendants of Vandals or Romans was found. It
is not yet clear whether the small percentage with African descent was in the region when the
Phoenicians arrived, or whether they arrived later. The second scenario fits the data better.

The Berber (and Tuareg) languages follow the Y-DNA characteristics and are the result of migrations
from Phoenicia.

Acknowledgements

I thank the people who tested their DNA and made it available in the datasets of yfull and
familytreedna. I also thank the discussion groups where interested persons and citizen scientists
contributed in questions and answers. I also thank Dr. Maarten Larmuseau, Yacine Kemouche, Ashraf
Taqatqa and Raf Ceulemans who gave replies to an earlier version of the document, which helped me
to improve the document.
Index
Abstract ................................................................................................................................................... 1
Introduction............................................................................................................................................. 1
Analysis of E-PF2546................................................................................................................................ 2
The M81 branches before and after E-PF2546 ....................................................................................... 2
M81 in the Maghreb................................................................................................................................ 3
Analysis of the non-E-M81 branch splits in the Magheb ........................................................................ 5
Analysis of CTS12227+PF6789 ................................................................................................................. 6
Analysis of all J1 branches in the Maghreb ............................................................................................. 6
Maghreb Y-DNA percentages and origin ................................................................................................. 8
Publications on ancient DNA in the line of E-M81 .................................................................................. 8
On the relation between ancient DNA of M81-branch in Ifri n’Amr or Moussa and the present M81-
descendants in the Maghreb................................................................................................................... 9
The yfull age normalization ................................................................................................................... 10
Female lines in Maghreb ....................................................................................................................... 12
Most likely history (Occam’s razor) ....................................................................................................... 12
Conclusion ............................................................................................................................................. 14
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