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In the overview of Greece (Fig. S41),Neolithic Greece, on the basis of available samples,
is composed of primarily Anatolian Neolithic-related ancestry as previously inferred.(9, 421)We
confirm the presence of some “eastern” CHG-related ancestry in the Peloponnese Neolithic,
234
although we infer that it was also present in the Neolithic of northern Greece(421), and also in
the Neolithic-to-Chalcolithic samples from Albania previously discussed. In the Peloponnese it
was accompanied by a small but significant (6±2%) amount of Levantine ancestry. The joint
presence of CHG/Levantine ancestry suggests that this non-Anatolian related ancestry may be
derived from a CHG/Levantine cline of which we have already seen the Mesopotamian Neolithic
was a part.
In the Bronze Age we group individuals into broad Minoan and Mycenaean groups which
are used as convenient labels to indicate Cretan and mainland Greece samples; we will look at
individual variation within these broad groups below. Anatolian Neolithic ancestry continues to
make up the greater part of the ancestry of the Bronze Age samples of both groups, but there is a
stronger presence of the “eastern” CHG and to lesser extent Levantine ancestry. This may reflect
a fresh pulse of “eastern” ancestry into the Aegean prior to the Bronze Age samples, as it in
Anatolia and the rest of West Asia that CHG ancestry exceeds the low levels that occurred in
southeastern Europe in the Neolithic. The proportion of eastern hunter-gatherer ancestry is very
small but is nevertheless different(16)in the combined Mycenaean sample at a significant 4±1%
with the Minoans at a non-significant 1±1%. The EHG ancestry is ~3-fold lower in the
Mycenaean samples than in Bronze Age samples from North Macedonia and Albania
immediately to the north of Greece and ~10-fold lower than in Moldova on the edge of the
steppe. Thus, our results suggest that although steppe-derived ancestry was present in Bronze
Age Greece it was quantitatively the weakest discernible component, only a little above the
practically non-existent Balkan hunter-gatherer ancestry.
The highest EHG ancestry is found in a previously published(16)low-coverage individual
from Armenoi in Crete at 24±6% but this is accompanied by a much smaller 3±7% of CHG
ancestry so it is difficult to interpret as being derived from steppe migrations where EHG/CHG
components are balanced. We do not overinterpret this Cretan outlier except to note it puts an
upper bound on the amount of EHG ancestry observed in the Bronze Age Aegean in a collective
sample of43 individuals. On the basis of this sample, it is justified to claim that while there was
variation in EHG ancestry in the Aegean, individuals of very high steppe ancestry such as the
ones found north of Greece were not common there during the Middle/LateBronze Age
time frame of ours samples.
Two individuals from Kastrouli near Delphi were from the Archaic age (I17962; 775-542
calBCE, and I17959 800-500 BCE). They were genetically similar to Mycenaean-era samples
from Kastrouli but their combined EHG ancestry was not significant (2±1%). They differed from
the Mycenaean sample set in having more Anatolian Neolithic and less Levantine-related
ancestry; their ancestral composition does not suggest any external influence between the Late
Bronze Age and Archaic periods as influence from the north would have introduced more
EHG/Balkan hunter-gatherer ancestry, and from the east more CHG/Levantine ancestry none of
which are observed for the Archaic individuals. Another individual from the vicinity of the
Palace of Nestor of Proto-Geometric/Early Iron Age time (I19368) has 14±5% which is similar
to the proportion seen north of Greece, albeit with a high standard error and a proportion that is
not significantly higher than the Mycenaean group as a whole. On the basis of these 3 post
Mycenaean individuals from Pylos and Kastrouli, it appears that some of the variation that
existed in the Mycenaean period persisted into the Iron Age without a sign of external influence.
Finally, we note that a Roman-era individual from Marathon (I7833; 252-392 calCE) is
within the range of ancestry for the population of the Bronze Age, although with somewhat more
eastern (CHG) ancestry; there would definitely have been an opportunity for such ancestry to
reach Greece in Hellenistic and Roman times, as evidenced also by the study of the population of
Rome in Republic and Imperial times.(436)On the basis of a single individual we cannot
conclude that there was a systematic Roman-era shift of the population in an eastward direction
as was proposed for Rome for Imperial times. Nonetheless, such “eastern”-shifted individuals
were described also in North Macedonia (above) and Croatia (below), and so may cumulatively
suggest that southeastern Europe also participated to some degree in the eastward shift of
ancestry that was also observed in Central Italy.
To better understand patterns of ancestry in the main Minoan and Mycenaean clusters, we
also plot finer-scale variation diagrams for these two groups(Fig. S42).
The Minoan individuals appear fairly homogeneous with few pairwise differences in
ancestry being significant. Two possible exceptions are a published individual(16)from Odigitria
(I9129) which is inferred to have more Anatolian Neolithic ancestry than average, and a newly
published Middle Minoan individual from Zakros (I14196) which has less Anatolian Neolithic
and more Levantine Neolithic ancestry than average.
Examining subsets of the Mycenaean group(Fig. S43)and focusing on the diagonal we
observe that most regional populations do not differ significantly from the population average in
the five components. Some notable exceptions are the inferred absence of EHG ancestry in a
low-coverage sample from ProskynasIV in Lokris (I6420_d; 1613-1509 calBCE), the less than
average Levantine ancestry in Attica with corresponding more Anatolian Neolithic ancestry, and
the corresponding 24±5% greater Anatolian Neolithic ancestry in Attica than in neighboring
Salamis. So,while there may have been some variation in terms of ancestry in the Mycenaean
world, this seems to be in terms of slightly different proportions of the major sources of ancestry.
We also looked at per-individual diagrams for the two populations with large sample sizes:
(Kastrouli, and the Palace of Nestor in Pylos)(Fig. S44)
The individuals from these two sites in the southwestern Peloponnese and central Greece do
not form site-specific clusters in terms of their ancestry. Some individuals differ in terms of their
ancestral composition from the overall mean or from each other, suggesting some level of
ancestral heterogeneity in Mycenaean Greece.
We highlight the case of the “Griffin Warrior(437)”,from the Palace of Nestor in Pylos
(I13519_d), a 30-35 year old male buried in a shaft grave with rich grave goods many of which
were produced in the Minoan world. The ancestral composition of this individual is
unremarkable suggesting that this elite individual did not belong to a genetically differentiated
population relative to the population of the Bronze Age Aegean at large, similar to another high
status individual from Peristeria(16)(I9033), thus showing no correlation between wealth and
ancestry. However, he is estimated as not having EHG ancestry and this is significantly less than
the Mycenaean population as a whole, thus lending some plausibility to potential Cretan
connections, although he is by no means the only Mycenaean individual to lack evidence of this
type of ancestry. In the PCA(6), the Griffin Warrior is near the center of the Bronze Aegean
cluster, in-between the Minoan and Mycenaean subclusters and appears to be (a) unremarkable
for a LBA individual from the Aegean, and (b) not clearly belonging to either of these two
groupings that form the structure of the Aegean population.
To test whether the Griffin Warrior was an outlier for the Palace of Nestor site in terms of
his EHG ancestry we computed the difference between each individual of this site and the
remaining individuals (Fig. S45). For the Griffin Warrior this difference is-5.9±1.1% which
corresponds to a 4.08e-8 probability that the difference is negative (which remains significant
even if we correct it for the 9 individuals considered). For most individuals the difference is not
significantly negative, while for I13510 and I13516 it is. Individual I13516 was buried just
outside the limits of the town surrounding the Palace of Nestor, while I13510 was buried in a
chamber tomb 500m from the Palace of Nestor. Thus, the individuals identified as having less
than average EHG ancestry in Pylos were buried in 3 different types of tombs associated with
different levels of prestige, and 3 different localities within the area
We also computed admixture proportions in our framework on 6 samples from Greece(31)
that were published during the course of our analysis and present them in together with other
comparative data(Fig. S46). Estimated admixture proportions for the Minoan sample from
Kephala Petras is within 1% of those of the rest of the Minoans. EHG ancestry of the Elati-
Logkas MBA samples from northern Greece is high (16±2%) and contrasts with that of the
Mycenaean population and reaches levels seen in neighboring Albania at Çinamak. The Logkas
samples are separated by a few hundred kilometers and a few centuries from the LBA
Mycenaeans of Central-Southern Greece and are an important datapoint in the Southeastern
European continuum between the low-EHG Mycenaean south and the higher-EHG north
PG 233
Supplementary Materials for
The genetic history of the Southern Arc: A bridge between West Asia and Europe
Iosif Lazaridis, Songül Alpaslan-Roodenberget al. 2022
file:///C:/Users/18509/Downloads/science.abm4247_sm.pdf
is composed of primarily Anatolian Neolithic-related ancestry as previously inferred.(9, 421)We
confirm the presence of some “eastern” CHG-related ancestry in the Peloponnese Neolithic,
234
although we infer that it was also present in the Neolithic of northern Greece(421), and also in
the Neolithic-to-Chalcolithic samples from Albania previously discussed. In the Peloponnese it
was accompanied by a small but significant (6±2%) amount of Levantine ancestry. The joint
presence of CHG/Levantine ancestry suggests that this non-Anatolian related ancestry may be
derived from a CHG/Levantine cline of which we have already seen the Mesopotamian Neolithic
was a part.
In the Bronze Age we group individuals into broad Minoan and Mycenaean groups which
are used as convenient labels to indicate Cretan and mainland Greece samples; we will look at
individual variation within these broad groups below. Anatolian Neolithic ancestry continues to
make up the greater part of the ancestry of the Bronze Age samples of both groups, but there is a
stronger presence of the “eastern” CHG and to lesser extent Levantine ancestry. This may reflect
a fresh pulse of “eastern” ancestry into the Aegean prior to the Bronze Age samples, as it in
Anatolia and the rest of West Asia that CHG ancestry exceeds the low levels that occurred in
southeastern Europe in the Neolithic. The proportion of eastern hunter-gatherer ancestry is very
small but is nevertheless different(16)in the combined Mycenaean sample at a significant 4±1%
with the Minoans at a non-significant 1±1%. The EHG ancestry is ~3-fold lower in the
Mycenaean samples than in Bronze Age samples from North Macedonia and Albania
immediately to the north of Greece and ~10-fold lower than in Moldova on the edge of the
steppe. Thus, our results suggest that although steppe-derived ancestry was present in Bronze
Age Greece it was quantitatively the weakest discernible component, only a little above the
practically non-existent Balkan hunter-gatherer ancestry.
The highest EHG ancestry is found in a previously published(16)low-coverage individual
from Armenoi in Crete at 24±6% but this is accompanied by a much smaller 3±7% of CHG
ancestry so it is difficult to interpret as being derived from steppe migrations where EHG/CHG
components are balanced. We do not overinterpret this Cretan outlier except to note it puts an
upper bound on the amount of EHG ancestry observed in the Bronze Age Aegean in a collective
sample of43 individuals. On the basis of this sample, it is justified to claim that while there was
variation in EHG ancestry in the Aegean, individuals of very high steppe ancestry such as the
ones found north of Greece were not common there during the Middle/LateBronze Age
time frame of ours samples.
Two individuals from Kastrouli near Delphi were from the Archaic age (I17962; 775-542
calBCE, and I17959 800-500 BCE). They were genetically similar to Mycenaean-era samples
from Kastrouli but their combined EHG ancestry was not significant (2±1%). They differed from
the Mycenaean sample set in having more Anatolian Neolithic and less Levantine-related
ancestry; their ancestral composition does not suggest any external influence between the Late
Bronze Age and Archaic periods as influence from the north would have introduced more
EHG/Balkan hunter-gatherer ancestry, and from the east more CHG/Levantine ancestry none of
which are observed for the Archaic individuals. Another individual from the vicinity of the
Palace of Nestor of Proto-Geometric/Early Iron Age time (I19368) has 14±5% which is similar
to the proportion seen north of Greece, albeit with a high standard error and a proportion that is
not significantly higher than the Mycenaean group as a whole. On the basis of these 3 post
Mycenaean individuals from Pylos and Kastrouli, it appears that some of the variation that
existed in the Mycenaean period persisted into the Iron Age without a sign of external influence.
Finally, we note that a Roman-era individual from Marathon (I7833; 252-392 calCE) is
within the range of ancestry for the population of the Bronze Age, although with somewhat more
eastern (CHG) ancestry; there would definitely have been an opportunity for such ancestry to
reach Greece in Hellenistic and Roman times, as evidenced also by the study of the population of
Rome in Republic and Imperial times.(436)On the basis of a single individual we cannot
conclude that there was a systematic Roman-era shift of the population in an eastward direction
as was proposed for Rome for Imperial times. Nonetheless, such “eastern”-shifted individuals
were described also in North Macedonia (above) and Croatia (below), and so may cumulatively
suggest that southeastern Europe also participated to some degree in the eastward shift of
ancestry that was also observed in Central Italy.
To better understand patterns of ancestry in the main Minoan and Mycenaean clusters, we
also plot finer-scale variation diagrams for these two groups(Fig. S42).
The Minoan individuals appear fairly homogeneous with few pairwise differences in
ancestry being significant. Two possible exceptions are a published individual(16)from Odigitria
(I9129) which is inferred to have more Anatolian Neolithic ancestry than average, and a newly
published Middle Minoan individual from Zakros (I14196) which has less Anatolian Neolithic
and more Levantine Neolithic ancestry than average.
Examining subsets of the Mycenaean group(Fig. S43)and focusing on the diagonal we
observe that most regional populations do not differ significantly from the population average in
the five components. Some notable exceptions are the inferred absence of EHG ancestry in a
low-coverage sample from ProskynasIV in Lokris (I6420_d; 1613-1509 calBCE), the less than
average Levantine ancestry in Attica with corresponding more Anatolian Neolithic ancestry, and
the corresponding 24±5% greater Anatolian Neolithic ancestry in Attica than in neighboring
Salamis. So,while there may have been some variation in terms of ancestry in the Mycenaean
world, this seems to be in terms of slightly different proportions of the major sources of ancestry.
We also looked at per-individual diagrams for the two populations with large sample sizes:
(Kastrouli, and the Palace of Nestor in Pylos)(Fig. S44)
The individuals from these two sites in the southwestern Peloponnese and central Greece do
not form site-specific clusters in terms of their ancestry. Some individuals differ in terms of their
ancestral composition from the overall mean or from each other, suggesting some level of
ancestral heterogeneity in Mycenaean Greece.
We highlight the case of the “Griffin Warrior(437)”,from the Palace of Nestor in Pylos
(I13519_d), a 30-35 year old male buried in a shaft grave with rich grave goods many of which
were produced in the Minoan world. The ancestral composition of this individual is
unremarkable suggesting that this elite individual did not belong to a genetically differentiated
population relative to the population of the Bronze Age Aegean at large, similar to another high
status individual from Peristeria(16)(I9033), thus showing no correlation between wealth and
ancestry. However, he is estimated as not having EHG ancestry and this is significantly less than
the Mycenaean population as a whole, thus lending some plausibility to potential Cretan
connections, although he is by no means the only Mycenaean individual to lack evidence of this
type of ancestry. In the PCA(6), the Griffin Warrior is near the center of the Bronze Aegean
cluster, in-between the Minoan and Mycenaean subclusters and appears to be (a) unremarkable
for a LBA individual from the Aegean, and (b) not clearly belonging to either of these two
groupings that form the structure of the Aegean population.
To test whether the Griffin Warrior was an outlier for the Palace of Nestor site in terms of
his EHG ancestry we computed the difference between each individual of this site and the
remaining individuals (Fig. S45). For the Griffin Warrior this difference is-5.9±1.1% which
corresponds to a 4.08e-8 probability that the difference is negative (which remains significant
even if we correct it for the 9 individuals considered). For most individuals the difference is not
significantly negative, while for I13510 and I13516 it is. Individual I13516 was buried just
outside the limits of the town surrounding the Palace of Nestor, while I13510 was buried in a
chamber tomb 500m from the Palace of Nestor. Thus, the individuals identified as having less
than average EHG ancestry in Pylos were buried in 3 different types of tombs associated with
different levels of prestige, and 3 different localities within the area
We also computed admixture proportions in our framework on 6 samples from Greece(31)
that were published during the course of our analysis and present them in together with other
comparative data(Fig. S46). Estimated admixture proportions for the Minoan sample from
Kephala Petras is within 1% of those of the rest of the Minoans. EHG ancestry of the Elati-
Logkas MBA samples from northern Greece is high (16±2%) and contrasts with that of the
Mycenaean population and reaches levels seen in neighboring Albania at Çinamak. The Logkas
samples are separated by a few hundred kilometers and a few centuries from the LBA
Mycenaeans of Central-Southern Greece and are an important datapoint in the Southeastern
European continuum between the low-EHG Mycenaean south and the higher-EHG north
PG 233
Supplementary Materials for
The genetic history of the Southern Arc: A bridge between West Asia and Europe
Iosif Lazaridis, Songül Alpaslan-Roodenberget al. 2022
file:///C:/Users/18509/Downloads/science.abm4247_sm.pdf