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Post by Admin on Aug 1, 2023 18:06:48 GMT
 Figure 2 Genetic differentiation of Bronze Age populations to present-day populations We plot the FST inbreeding coefficient (Methods) between newly reported populations and present-day West Eurasian populations which shows a pattern of genetic affinity between Bronze Age and present-day populations from the corresponding broad geographical regions. (a) Mycenaeans, (b) Minoans from Hagios Charalambos (Lasithi regional unit), (c) Minoans from Moni Odigitria (Heraklion regional unit), (d) southwestern Bronze Age Anatolians. The same pattern also applies to Bronze Age populations from other regions of West Eurasia (Extended Data Fig. 5). The Minoans and Mycenaeans, sampled from different sites in Crete and mainland Greece, were homogeneous, supporting the genetic coherency of these two groups. Differences between them are only relative, viewed against their broad overall similarity to each other and to the southwestern Anatolians, sharing in both the ‘local’ Anatolian Neolithic-like farmer ancestry and the ‘eastern’ Caucasus-related admixture. Two key questions remain to be addressed by future studies. First, when did the common ‘eastern’ ancestry of both Minoans and Mycenaeans arrive in the Aegean? Second, is the ‘northern’ ancestry in Mycenaeans due to sporadic infiltration of Greece, or the result of a rapid migration as in Central Europe6? Such a migration would support the idea that Proto-Greek speakers29 formed the southern wing of a steppe intrusion of Indo-European speakers. Yet, the absence of ‘northern’ ancestry in the Bronze Age samples from Pisidia, where Indo-European languages were attested in antiquity, casts doubt on this genetic-linguistic association, with further sampling of ancient Anatolian speakers needed. Whatever the answer to these questions, the discovery of at least two migration events into the Aegean in addition to the first farming dispersal and before the Bronze Age, and of additional population change since that time, supports the view that the Greeks did not emerge fully-formed from the depths of prehistory, but were, indeed, a people ‘ever in the process of becoming.’30 |
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Post by Admin on Aug 2, 2023 20:12:34 GMT
Extended Data Figure 1  ADMIXTURE analysis ADMIXTURE analysis with K=2 to K=17 is shown. 351 ancient and 2,616 present-day individuals were used in this analysis; ancient samples and present-day Greeks are displayed. To avoid visual clutter of labels, individuals in populations with sample size ≤5 are shown with thicker lines. Extended Data Figure 2  Symmetry testing of Aegean Bronze Age populations The statistic f4(X, Y; Test, Chimp) is shown with ±3 standard errors. Each panel is titled with the pair X, Y. Populations are ordered according to the value of the statistic. Positive values indicate that Test shares more alleles with X than Y and negative values that it shares more with Y than X. (a) ‘northern’ and ‘eastern’ populations share more alleles with Minoans than with Neolithic Greece. (b) ‘northern’ and ‘eastern’ populations share more alleles with Mycenaeans than with Neolithic Greece. (c) Minoans from Lasithi and Moni Odigitria are symmetrically related to diverse populations. (d) Neolithic populations from Anatolia, Europe, Greece, and the Levant share fewer alleles with Mycenaeans than with Minoans. Extended Data Figure 3  Symmetry testing of Anatolian Bronze Age populations The statistic f4(X, Y; Test, Chimp) is shown with ±3 standard errors. Each panel is titled with the pair X, Y. Populations are ordered according to the value of the statistic. Positive values indicate that Test shares more alleles with X than Y and negative values that it shares more with Y than X. (a) European, Siberian, and Caucasus hunter-gatherers share fewer alleles with Bronze Age Anatolians from Harmanören Göndürle than with a Chalcolithic Anatolian from Barcın. (b) Bronze Age Anatolians differ from Neolithic ones in sharing more alleles with populations of Iran, the Caucasus, and the Steppe than with those of Europe. (c) Bronze Age Anatolians differ from Minoans in sharing more alleles with populations from Neolithic Iran than Neolithic Anatolia and Europe. (d) Bronze Age Anatolians differ from Mycenaeans in sharing more alleles with Neolithic and Bronze Age populations of the Levant. |
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Post by Admin on Aug 3, 2023 18:53:11 GMT
Extended Data Figure 4  f3-statistics of Mycenaeans as a target with different pairs of reference populations We show the value of the statistic f3(Ref1, Ref2; Mycenaean) and ±3 standard errors; only the population pairs (Ref1, Ref2) for which the Z-score of the statistic is <−2 are shown. Negative values indicate that the Mycenaean population is admixed from sources related to the two reference populations. Extended Data Figure 5  Correspondence of qpAdm estimates with PCA As a way to validate qpAdm models of admixture for Myceneans from three ancestral populations (Anatolia_N or Minoan_Lasithi), (Armenia_ChL or Armenia_MLBA), (Steppe_EMBA, Steppe_MLBA, Europe_LNBA), representing substratum, ‘eastern’, and ‘northern’ ancestry respectively (Supplementary Information, section 2), we plot the qpAdm-predicted position in the PCA space of Fig. 1 vs. the actual position of the Mycenaean population. Extended Data Figure 6  Comparison of Mycenaeans and simulated admixed populations We simulate admixed individuals with known ancestry from three ancestral populations (Anatolia_N or Minoan_Lasithi), (Armenia_ChL or Armenia_MLBA), (Steppe_EMBA, Steppe_MLBA, Europe_LNBA), representing substratum, ‘eastern’, and ‘northern’ ancestry respectively (Methods; Supplementary Information, section 2). The maximum |Z|-score of statistics f4(Mycenaean, Simulated; Outgroup1, Outgroup2) is plotted with circles of varying size (proportional to log|Z|) for each assignment of ancestry proportions. The best estimate (red) corresponds to the proportions that minimize |Z|, and they are compared against the qpAdm estimate for the same ancestral sources (blue). |
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Post by Admin on Aug 6, 2023 19:47:41 GMT
Extended Data Figure 7  FST between Bronze Age and present-day West Eurasian populations (a) The population of Early Bronze Age Armenia4 shows an affinity to present-day populations from Armenia, Anatolia, the Caucasus, and Iran, as does (b) Middle/Late Bronze Age Armenia4,9. (c) The Bronze Age Levant4 has an affinity to Levantine and Arabian populations. (d) Late Neolithic/Bronze Age Europeans1,6,9,43 most resemble present-day northern/central Europeans, as do (e) Early/Middle Bronze Age steppe populations1,6,9, who also resemble populations of the northeast Caucasus, while (f) Middle/Late Bronze Age steppe populations resemble central/northern Europeans1,9. Jewish populations are plotted with a square to distinguish them from non-Jewish populations from the same geographical area. The plots for the newly reported populations of Mycenaeans, Minoans, and Bronze Age Anatolians are shown in Fig. 2. Extended Data Figure 8  Symmetry testing of Mycenaeans with Modern Greek populations The statistic f4(Mycenaean, Modern Greek; Test, Chimp) is shown with ±3 standard errors. Modern Greeks share fewer alleles with Levantine/Anatolian/European Neolithic populations and with Minoans than Mycenaeans do, suggesting a dilution of early Neolithic ancestry since the Bronze Age. Human Origins genotype data: (a) Greeks from the Coriell repository10, (b) Greeks from Thessaloniki10, (c) Cypriots10. Whole genome data: (d) Cretans40. Illumina genotype data: (e) Greeks from Thessaly41, (f) Greeks from Central Greece41, (g) Greeks from the study by Hellenthal et al.27
Extended Data Table 1
Information on ancient samples reported in this study
Dates marked simply as BCE are based on the associated archaeology of the samples. Dates marked as calBCE are based on radiocarbon dating of the samples (Supplementary Information, section 1).
Individual_ID Genotype_ID Other_ID Source Date Population_Label Location Country Latitude Longitude Sex Coverage Autosomal_SNPs mtDNA Y-chromosome
I2937 I2937 A2197 1240K 5419±41
cal BC Greece_N Diros,
Alepotrypa
Cave Greece 36.64 22.38 F 0.870 481848 K1a26
I0071 I0071 Lasithi4 1240K 2000-1700
BCE Minoan_Lasithi Hagios
Charalambos
Cave,
Lasithi,
Crete Greece 35.08 25.83 F 7.312 953157 U5a1
I0070 I0070 Lasithi2 1240K 2000-1700
BCE Minoan_Lasithi Hagios
Charalambos
Cave,
Lasithi,
Crete Greece 35.08 25.83 M 1.267 619767 H13a1 J2a1d
I0073 I0073 Lasithi7 1240K 2000-1700
BCE Minoan_Lasithi Hagios
Charalambos
Cave,
Lasithi,
Crete Greece 35.08 25.83 M 1.481 643360 H J2a1
I0074 I0074 Lasithi9 1240K 2000-1700
BCE Minoan_Lasithi Hagios
Charalambos
Cave,
Lasithi,
Crete Greece 35.08 25.83 F 0.874 506434 H5
I9005 I9005 Lasithi17 1240K 2000-1700
BCE Minoan_Lasithi Hagios
Charalambos
Cave,
Lasithi,
Crete Greece 35.08 25.83 F 1.351 388859 H
I9006 I9006 Salamis31 1240K 1411-1262
cal BCE
(3067 ± 25 BP,
DEM-2905) Mycenaean Agia
Kyriaki,
Salamis Greece 37.97 23.50 F 1.387 361193 X2d
I9123 I9123 S-EVA 1263
Armenoi 503 1240K 1370-1340 BCE Crete_Armenoi Armenoi,
Crete Greece 35.45 24.17 F 0.041 45158 U5a1
I9127 I9127 12V t2 1240K 2900-1900
BCE Minoan_Odigitria Moni
Odigitria,
Heraklion,
Crete Greece 35.05 24.81 F 0.035 36475 J2b1a1
I9128 I9128 13V t2 1240K 2900-1900
BCE Minoan_Odigitria Moni
Odigitria,
Heraklion,
Crete Greece 35.05 24.81 F 0.016 17081 I5
I9129 I9129 14V t2 1240K 2900-1900
BCE Minoan_Odigitria Moni
Odigitria,
Heraklion,
Crete Greece 35.05 24.81 F 0.063 63986 H+163
I9130 I9130 16V Tholos 1240K 2900-1900
BCE Minoan_Odigitria Moni
Odigitria,
Heraklion,
Crete Greece 35.05 24.81 M 0.086 92186 U3b3 G2a2b2
I9131 I9131 19V t2 1240K 2900-1900
BCE Minoan_Odigitria Moni
Odigitria,
Heraklion,
Crete Greece 35.05 24.81 F 0.095 96946 K1a2
I9010 I9010 Galatas19 1240K 1700-1200
BCE Mycenaean Galatas
Apatheia,
Peloponnese Greece 37.50 23.45 F 0.379 242265 X2
I9033 I9033 Peristeria4 1240K 1416-1280
cal BCE
(3084 ± 24 BP,
DEM-2903) Mycenaean Peristeria
Tryfilia,
Peloponnese Greece 36.92 21.70 F 0.439 248912 H
I9041 I9041 Galatas4 1240K 1700-1200
BCE Mycenaean Galatas
Apatheia,
Peloponnese Greece 37.50 23.45 M 1.558 417898 X2 J2a1
I2495 I2495 A4-1 1240K 2558-2295
caIBCE
(3925±35 BP,
Poz-81111) Anatolia_BA Harmanӧren-
Gӧndürle
Hӧyük,
Isparta Turkey 37.92 30.71 M 1.981 637146 H J1a
I2499 I2499 UC1 1240K 2836-2472
caIBCE
(4040±35 BP,
Poz-82213) Anatolia_BA Harmanӧren-
Gӧndürle
Hӧyük,
Isparta Turkey 37.92 30.71 F 0.285 243348 K1a2
I2683 I2683 G3-95 1240K 2500-1800
BCE Anatolia_BA Harmanӧren-
Gӧndürle
Hӧyük,
Isparta Turkey 37.92 30.71 F 3.695 749308 T2b
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Post by Admin on Aug 10, 2023 19:23:48 GMT
Phenotypic inference of ancient individuals We list the probability assignments for different phenotypes by HIrisPlex26 and an assessment of the phenotype. We generate 100 random replicates of the genotypes of each individual, listing the standard deviation in parentheses (Supplementary Information, section 4).
ID Population PBlueEye PIntermediateEye PBrownEye PBlondHair PBrownHair PRedHair PBlackHair PLightHair PDarkHair Hair Color Eye Clor
I2495 Anatolia_BA 1.6 (4.4) 3.6 (3.9) 94.9 (8.3) 10.7 (6.1) 51.6 (6.4) 0.1 (0.1) 37.6 (9.3) 18.0 (11.7) 82.0 (11.7) Brown Brown
I2499 Anatolia_BA 16.6 (28.3) 7.4 (2.2) 76.0 (28.7) 2.2 (2.2) 64.7 (11.8) 2.0 (5.3) 31.1 (13.8) 12.9 (20.1) 87.1 (20.1) Brown Blue or Brown
I2683 Anatolia_BA 0.3 (0.9) 1.3 (1.7) 98.4 (2.6) 3.3 (2.5) 33.0 (4.6) 0.0 (0.0) 63.7 (7.0) 4.9 (4.5) 95.1 (4.5) Black Brown
I2937 Greece_N 0.3 (1.3) 2.2 (1.9) 97.5 (3.2) 3.6 (1.9) 33.9 (6.2) 0.1 (0.0) 62.4 (7.4) 6.7 (4.3) 93.3 (4.3) Black Brown
I0070 Minoan_Lasithi 0.4 (1.8) 2.2 (1.9) 97.4 (3.7) 30.4 (5.1) 66.4 (5.9) 3.2 (0.9) 0.0 (0.0) 100.0 (0.0) 0.0 (0.0) Brown Brown
I0071 Minoan_Lasithi 0.0 (0.0) 0.2 (0.0) 99.8 (0.0) 0.4 (0.0) 20.3 (0.0) 0.0 (0.0) 79.3 (0.0) 0.5 (0.0) 99.5 (0.0) Black Brown
I0073 Minoan_Lasithi 0.1 (0.7) 1.7 (1.4) 98.2 (2.2) 12.5 (3.4) 61.1 (1.2) 0.2 (0.1) 26.2 (2.7) 32.4 (8.8) 67.6 (8.8) Brown Brown
I0074 Minoan_Lasithi 0.0 (0.0) 1.3 (0.3) 98.7 (0.4) 9.3 (3.2) 54.8 (8.5) 0.1 (0.1) 35.8 (10.5) 18.8 (10.3) 81.2 (10.3) Brown Brown
I9005 Minoan_Lasithi 5.2 (0.0) 11.6 (0.0) 83.2 (0.0) 49.6 (1.4) 38.8 (1.2) 4.2 (0.5) 7.4 (0.7) 85.6 (1.7) 14.4 (1.7) Blond or Brown Brown
I9006 Mycenaean 0.0 (0.0) 1.1 (0.4) 98.9 (0.4) 8.7 (4.9) 59.9 (6.4) 1.8 (2.9) 29.6 (11.8) 25.7 (16.5) 74.3 (16.5) Brown Brown
I9033 Mycenaean 0.4 (1.0) 1.6 (1.9) 98.0 (3.0) 4.6 (3.9) 51.0 (6.3) 0.1 (0.5) 44.2 (9.8) 10.5 (13.2) 89.5 (13.2) Brown Brown
I9041 Mycenaean 1.4 (0.5) 5.3 (1.0) 93.3 (1.4) 7.8 (0.7) 63.2 (2.0) 0.2 (0.4) 28.7 (2.3) 21.2 (2.5) 78.8 (2.5) Brown Brown
The origin of the Mycenaeans has been intensely debated. Various theories have been
proposed to respond to the question of if, when and in what circumstances Greek speakers or
their linguistic ancestors, speaking a language that later developed into Greek, entered the
Aegean. One theory attributes the origin of Greek speakers to the Balkans, from which waves
of Indo-European speakers flowed into the north of Greece during the Bronze Age. These
people came from the Eurasian steppe north of the Black and Caspian seas, [9] and they are
referred to as the Proto Indo-Europeans. These migrants, together with the local population
they encountered, then combined to form the ancestors of the Mycenaeans and later Greek
speakers. [10-12] One problem with this theory is that the material culture relationship of
Bronze Age populations of the Aegean with populations far to the north is very tenuous. [11-12]
Another theory traces the origin of the Proto-Greeks further back in time, to approximately
3000 BCE at the start of the Early Bronze Age. [13] It proposes that as migrants, they filled a
largely depopulated landscape.
An additional hypothesis for the origin of the Greeks goes even further back to the seventh
millennium BCE and is associated with the view that the Greeks are descended from the first
farmers who migrated into Europe from Anatolia. Alternatively, a very late origin of
Mycenaean elites, associated with chariot riding warriors from the Caucasus in approximately
1600 BCE and characterised by those buried in the Shaft Graves at Mycenae, has also been proposed. [14]
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