The diverse genetic origins of a Classical period Greek army

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By studying genome-wide data from 54 individuals from eighth- to fifth-century Sicily, we gain insights into the composition of Classical Greek armies (ca. fifth c. BCE) and the populace of a Greek colony. The presence of mercenaries in Greek armies fighting in the Mediterranean, as early as 480 BCE and with origins as far away as northern Europe and the Caucasus, is absent from historical texts and thus so far underappreciated in ancient classical scholarship. Our interdisciplinary study both underlines the value of integrating genetic studies to complement archaeological and historical research and highlights the importance of warfare in facilitating continental-scale human mobility, cultural contact, and cooperation in the Mediterranean of the Classical period.




Abstract


Trade and colonization caused an unprecedented increase in Mediterranean human mobility in the first millennium BCE. Often seen as a dividing force, warfare is in fact another catalyst of culture contact. We provide insight into the demographic dynamics of ancient warfare by reporting genome-wide data from fifth-century soldiers who fought for the army of the Greek Sicilian colony of Himera, along with representatives of the civilian population, nearby indigenous settlements, and 96 present-day individuals from Italy and Greece. Unlike the rest of the sample, many soldiers had ancestral origins in northern Europe, the Steppe, and the Caucasus. Integrating genetic, archaeological, isotopic, and historical data, these results illustrate the significant role mercenaries played in ancient Greek armies and highlight how participation in war contributed to continental-scale human mobility in the Classical world.



https://www.pnas.org/doi/10.1073/pnas.2205272119
 
"The Civilian Population of Himera and Surrounding Regions.

We find that the IA Sicilians (Sicily_IA), associated with the Sicani culture, form a homogenous cluster distinct from most of the individuals excavated at Himera. They fall in the PC space occupied by earlier Middle and Late Bronze Age Sicilians (Sicily_MBA and Sicily_LBA; Fig. 2A) and display similar and homogenous proportions of the genetic components maximized in WHGs, CHGs, and EEFs in the ADMIXTURE analysis (blue, green, and orange in Fig. 2B, respectively). Using qpAdm, this group can be modeled as an admixture of four sources that distantly contributed to the genetic composition of Europeans (P = 0.179): Northwestern Anatolian Neolithic farmers (Turkey_Barcin_N; 76.4 ± 1.2%), WHGs (6.4 ± 1.0%), early farmers from Iran (Iran_GanjDareh_N; 6.3 ± 1.5%), and Early Bronze Age (EBA) Steppe herders associated with the Yamnaya cultural complex (Russia_Samara_EBA_Yamnaya; 10.9 ± 1.6%), which indicates an increase of Iranian-related admixture compared with the preceding LBA Sicily group, which can be best modeled without that component (SI Appendix, Table S8). Models using more proximal Bronze Age (BA) to Iron Age (IA) ancestry sources are not rejected for two-way mixtures between a local Sicilian source (Sicily_LBA, Sicily_LBA_I10371, or Sicily_MBA) on the one hand (57.1–98.7%) and Italy_IA_Republic.SG, Spain_BA, Spain_IA, Armenia_LBA.SG, or Balkan_IA (1.3–42.9%) on the other hand (SI Appendix, Table S9). This indicates that the population might not have been completely continuous to the BA inhabitants of Sicily sampled to date and instead received some gene flow from other populations that are most plausibly from outside of Sicily.
We caution that there are only published data from 15 individuals from BA Sicily, leaving the possibility that some unsampled Sicilian populations were genetically more continuous with later groups. The potential connection to Spain_IA is intriguing, as structural similarities in EBA fortifications have been documented at Castelluccio near Syracuse and artifacts, such as Bell Beakers, have been found at BA sites near Palermo (51). The arrival of amber on Sicily from Iberia has been documented to at least the third century BCE (52), if not earlier (53). Additionally, laterally spiked axes have been discovered on Sicily that likely derive from Iberian metallurgical traditions, suggesting wide Mediterranean trade networks (54). However, Leighton (1999) notes that, although “it has often been supposed that Sicilian beakers derive from Spanish types…there is no sure evidence that they represent actual imports, and since North Italian finds have multiplied in recent years and a central European origin for the form now seems likely, the case for a direct Iberian link is not as compelling as it was” (9). Ancient historians, who plausibly had access to sources that have since been lost, also discussed the possibility of connections to Iberia: Thucydides wrote of the Sicani that they were not indigenous to the island but settlers from Iberia (Thuc. 6.2.2) (55), but Diodorus Siculus disagreed with this claim and instead name them as the original inhabitants of Sicily (Diod. 5.2.4). Of course, the accounts of ancient historians writing hundreds or even thousands of years after the events they describe should not be taken as direct evidence but only as a representation of even older oral history, which is important to consider but may not be based on fact at all. Our results offer tentative support to both scenarios; however, it is unlikely that the Sicani were descended only from Iberian or other non-Sicilian populations. One line of evidence for some local genetic continuity is the almost exclusive presence of Y-chromosomal haplogroup G-Z1903 and its derivates among the males (Dataset S6), a lineage already found among Sicilian inhabitants in the Middle Bronze Age (MBA) and LBA (12) and otherwise unreported from any pre- or early historic contexts, including Iberia. One male belongs to haplogroup R-FT40455, which is a specific subtype of haplogroup R-DF27, which could reflect an Iberian source of ancestry, as it is much more common in Iberia from the BA onward than in any other region (19, 56). R-DF27 has also been observed in individuals of the EBA in Sicily (12) and thus could have that persisted to the time of the Sicani culture. We find overall low levels of runs of homozygosity among the Sicily_IA and preceding BA individuals (SI Appendix, Fig. S14 and Table S18), providing no evidence for endogamous practices and pointing toward large effective population sizes (57)....."
 
"Diverse Ancestry among Himera’s Soldiers of the 480 BCE Battle.
Most Himerans associated with the battles can be found clustering on the PCA closely with individuals from the Greece_LBA, consistent with a major contribution of individuals of primarily Greek ancestry in the Himeran forces and substantial genetic continuity between the LBA period in Greece and fifth-century-BCE Greek colonies in Sicily. These soldiers with at least some Greek ancestry could have been inhabitants of the colony or supporting armies from other colonies, such as Syracuse. Seven of the 16 soldiers of the 480 BCE battle (Sicily_Himera_480BCE_1) and all 5 of the soldiers of the 409 BCE battle (Sicily_Himera_409BCE) are part of this main genetic cluster. Using the qpWave/qpAdm framework, we can model each of the soldiers in these two groups as deriving their ancestry either 100% from a group related to Greece_LBA or from an admixture between a Sicilian LBA or IA source and an Aegean-related source in varying proportions (SI Appendix, Tables S16 and S17), suggesting that many soldiers (and all studied from the 409 BCE battle) were plausibly the descendants of the Greek colonizers of Sicily and that intermarriage between Greeks and Sicilian locals was practiced (63). This genetic evidence adds to our understanding of social practices in Greek colonies in Sicily, because it is difficult to detect intermarriage archaeologically due to the fact that the presence of local Sicilian objects at Greek colonies may represent trade rather than intermarriage (59, 64, 65). Genetic data also provide information that may be invisible archaeologically, because individuals would have had the agency to decide which burial traditions and objects to adopt when interring their dead, and these archaeological identity markers, and the way individuals viewed their own identities, may or may not align with genetic ancestry (59, 66).


Beyond the intermarriage between locals and Himera’s Greek settlers, it is important to consider the possibility that the composition of the population might also have been influenced by a large-scale influx of Dorians from Agrigento after a political takeover by the Agrigentine tyrant, Theron, in 476 BCE, during which thousands of Himera’s inhabitants were killed (Diod. 11.49); however, the limited sample size available for analysis at this point and the lack of comparative data from Dorian contexts does not allow us to test this hypothesis. In any case, both groups of soldiers are heterogeneous in their admixture with local Sicilians, indicating a different genetic history for each individual; while we can compellingly document that significant genetic variation existed, a larger sample size than we have available would be necessary to adequately characterize the distribution of genetic variation encompassed by Sicilian Greek ethnicity in this period.
Among the soldiers of the 480 BCE battle, we find nine individuals that carry genetic ancestry not consistent with the first group (Fig. 2 and SI Appendix, Figs. S10–S13). We tested ancestry models with qpAdm for the higher coverage individuals.
Sicily_Himera_480BCE_2 consists of two outlying individuals (I10946/W1771 and I10950/W814) that fall on the PCA intermediate between the main cluster and central European individuals, a differentiation also indicated by the relatively higher proportion of the WHG genetic cluster in ADMIXTURE. Testing a wide range of possible BA and IA sources with qpAdm, valid models of ancestry consist of mixtures of one source related to central or eastern Mediterranean groups (Sicilian, Aegean, or Balkan) and one source related to central or western European groups (France, Spain, Czechia, or Hungary). A genetic origin in the Balkans is suggested by their Y chromosomes, belonging to the E-V13 lineage, which has its highest modern-day frequency in that region (67)."

 
"Two individuals (I10943/W0396 and I10949/W0403; Sicily_Himera_480BCE_3) fall with modern northeastern European groups and eastern Baltic populations of the first millennium BCE and can be modeled using exclusively BA individuals from Lithuania as a proxy source (P = 0.129).
One low-coverage individual, I17870/W0336, falls intermediate between Sicily_Himera_480BCE_2 and Sicily_Himera_480BCE_3 on both PCA and with respect to the main ancestry clusters inferred from ADMIXTURE (Fig. 2).

Two (I10944/W0461 and I10947/W1774; Sicily_Himera_480BCE_4) fall with individuals from IA nomadic contexts in the Eurasian Steppe and carry in the ADMIXTURE analysis genetic components maximized in Han and Karitiana (Native Americans) that characterize most IA Steppe nomads (light green and purple, respectively, in SI Appendix, Fig. S2B). In qpAdm, their ancestry is consistent, with around 85–89% deriving from IA Central steppe nomads and 11–15% from an Aegean-like source, an admixture that plausibly could have taken place among the genetically diverse populations of the Steppe (68, 69). Their mitochondrial DNA (mtDNA) haplogroups suggest east Eurasian genetic roots: A6a, found so far only in modern-day China (70, 71), and N1a1a1a, restricted to Russia, Kazakhstan, and Mongolia (72)."





"We analyzed 11 individuals, believed to represent civilians, from single graves in Himera’s West necropolis, which was in use during the sixth to fifth centuries BCE and potentially spanning the entire occupation of the site, as analyses of burial inclusions to identify temporal information for individuals at Himera is still ongoing. Only three individuals have sufficient coverage (at least 50,000 SNPs covered) to perform formal statistics, but all individuals are consistent with a broadly Central or Eastern Mediterranean genetic profile in PC analysis (PCA) and ADMIXTURE (Fig. 2 and SI Appendix, Figs. S9–S12). Of the three higher-coverage individuals, both I20166/W3182 and I20168/W3702 are cladal with Greek_LBA (P = 3.55 × 10−2 and 7.80 × 10−2, respectively). I20163/W1838 appears distinct, with models of ancestry with the highest P values (P > 0.1) involving a contribution of 38–55% of his ancestry from a local Sicilian MBA, LBA, or IA source and around 45–62% from a group closely related to Punic individuals from Sardinia that harbor North African ancestry or alternatively around 87.6 ± 3.1% Sicily_IA ancestry with 12.4± 3.1% ancestry deriving from a group represented by genetic outlier from Chalcolithic Sardinia carrying fully North African ancestry (SI Appendix, Table S14). It is notable that four of the eight individuals that could be analyzed with ADMIXTURE carry a small proportion (2.2–3.8%) of a genetic component maximized in Africans that is not found to the same extent in our other Sicilian individuals but appears in Punic and Levantine individuals (Fig. 2B), and the same individuals are shifted toward Levantine groups on the worldwide PCA (SI Appendix, Fig. S11B). Considering the low coverage of the civilian individuals, comparisons of within-group variance and centroid distance on the first two PCs nevertheless suggest that the civilian sample overlapped with the soldiers of both battles in the groups Sicily_Himera_480BCE_1 and Sicily_Himera_409BCE, described below (P = 0.218, one-way PERMANOVA (58) of centroid distance). The Himera civilian individuals were, however, genetically significantly more diverse (P = 2.51–05, t test on Euclidean distance to the group centroid) than the soldiers, who derived most of their ancestry from Aegean-related populations and plausibly represented Greek settlers or their descendants. This greater genetic diversity is likely not simply explained by the lower coverage of the civilian individuals (SI Appendix, Figs. S12 and S15). A greater sample size ideally from a wider diversity of burial contexts at Himera would be necessary to test whether they could represent a genetically stratified subset of a more-diverse civilian population. Greek colonies were the meeting grounds of culturally and genetically diverse people (16, 17, 59). Those living in Greek poleis and apoikiai (independent cities and colonies, respectively) included people with ancestors from the Greek mainland and Greek isles but also people from the colony’s hinterlands and members of other cultural groups, such as Phoenicians and Etruscans, which could explain the signal we observe (2, 59–61). Himera, especially, may have had strong Phoenician ties due to its proximity to neighboring Punic settlements and the Himeran tyrant Terillus’ alliance with Carthage (Hdt. 7.165).
The individual buried at Himera’s East necropolis, which was in use from the seventh to fifth centuries, directly attests to the incorporation of local people into the populace of the colony, as this person is best modeled with qpAdm as deriving his ancestry from the contemporaneous Sicilian_IA group (P = 0.695). Potential social stratification has been suggested associated with the burial sites, as individuals interred in the East necropolis show lower δ13C and δ15N values and higher prevalence of skeletal pathology than those at the West necropolis, indicating a different access to resources (62). However, due to the limited number of samples, we cannot test whether such a significant stratification existed with respect to genetic ancestry.

Finally, one outlier (I10951/W0653; Sicily_Himera_480BCE_5) falls with modern Caucasus populations and intermediate to ancient Steppe and Caucasus individuals on the PCA and carries the highest proportion of the CHG component in ADMIXTURE (Fig. 2). A single one-way model with a group closely related to Armenia_MBA as the source fit the data (P = 0.293). Similarly, the second low-coverage individual, I17872/W0428, falls closest to populations from the Caucasus on the PCA (Fig. 2A).
Outgroup-f3 statistics with ancient groups reflect the same general patterns of genetic affinities (SI Appendix, Fig. S16), as do proportions of distal ancestry estimated with qpAdm (SI Appendix, Table S8 and Dataset S5). The groups can all be modeled as deriving differing proportions of ancestry from groups related to Turkey_N_Barcin, WHG, Iran_GanjDareh_N/CHG, and Russia_Samara_EBA_Yamnaya, except for group 4, which requires the addition of Siberian ancestry."
 
Maybe first occurrence of E-V13 in ancient Greek site?


Sicily_Himera_480BCE_lc I17872 E1b1b1a1b1 E-V13 Excluded sister/downstream haplogroups:
E-BY5026,E-FT146201,E-Y81971,E-Y92017,E-BY20074,E-Y138701,E-Z38770,E-BY5814,E-A779
Sicily_Himera_480BCE_2 I10946 E1b1b1a1b1a15a~ Excluded sister/downstream haplogroups: E-BY6357* E-BY6357+:BY6357(15728070G>T:T)
Sicily_Himera_480BCE_2 I10950 E1b1b1a1b1a6a~ Excluded sister/downstream haplogroups : E-CTS6377* E-BY95428,E-CTS9320



Sicily_Himera_480BCE_2 consists of two outlying individuals (I10946/W1771 and I10950/W814) that fall on the PCA intermediate between the main cluster and central European individuals, a differentiation also indicated by the relatively higher proportion of the WHG genetic cluster in ADMIXTURE. Testing a wide range of possible BA and IA sources with qpAdm, valid models of ancestry consist of mixtures of one source related to central or eastern Mediterranean groups (Sicilian, Aegean, or Balkan) and one source related to central or western European groups (France, Spain, Czechia, or Hungary). A genetic origin in the Balkans is suggested by their Y chromosomes, belonging to the E-V13 lineage, which has its highest modern-day frequency in that region (67).

 
It is first proof that e-v13 was
Present in ancient greeks to some extent

Even though it was missing from previews papers

P.s
By the way [I]The N case in 480 bc himera is mindblowing same goes for r1a cases [/I]
Would be happy to see the carthegenian army
Y haplogroups also maybe there was e-m81 among them but i don't see data about them
In this paper bummer
 
It is first proof that e-v13 was
Present in ancient greeks to some extent

Even though it was missing from previews papers
P.s
By the way [I]The N case in 480 bc himera is mindblowing same goes for r1a cases [/I]
Would be happy to see the carthegenian army
Y haplogroups also maybe there was e-m81 among them but i don't see data about them
In this paper bummer
E-V13 was kinda expected in classical era Greeks. N is indeed mindblowing. Seima-Turbino descendant Scythian or Thracian?
 
There's a neat PCA at page 38 Appendix 01 (PDF). None of the Greeks proper are E-v13
 
It is first proof that e-v13 was
Present in ancient greeks to some extent

Even though it was missing from previews papers
P.s
By the way [I]The N case in 480 bc himera is mindblowing same goes for r1a cases [/I]
Would be happy to see the carthegenian army
Y haplogroups also maybe there was e-m81 among them but i don't see data about them
In this paper bummer

It's an outlier though.
 
There's a neat PCA at page 38 Appendix 01 (PDF). None of the Greeks proper are E-v13
Yeah even in Hellenistic Anatolia the E-V13 line is a Thracian subclade. One would think more E-V13 would pop out in Greek colonies.
 
It is first proof that e-v13 was
Present in ancient greeks to some extent

Even though it was missing from previews papers
P.s
By the way [I]The N case in 480 bc himera is mindblowing same goes for r1a cases [/I]
Would be happy to see the carthegenian army
Y haplogroups also maybe there was e-m81 among them but i don't see data about them
In this paper bummer

According to the paper this sample does not belong to the Greek cluster and it has Balkan origin.
 
According to the paper this sample does not belong to the Greek cluster and it has Balkan origin.
But Thracians are in the Greek cluster? I wonder what they mean with "Balkans"?
 
Lazaridis in twiter

"There are many interesting findings, but I will highlight a few:First, many of the 5th c. BC Greek combatants were genetically very similar to Bronze Age Mycenaean Greeks (a thousand years earlier).

Thus, we may conclude that the Ionian and Dorian ethnē -from which the Sicilian Himerans were drawn, according to historical sources- did not represent the migration of a genetically dissimilar population into post-Mycenaean Greece.

Surprisingly, men of diverse genetic background (neither Greek nor Sicanian) were buried in mass graves in the city's West Necropolis: these men were drawn from the east Baltic, the Eurasian steppe, the Balkans, and the Caucasus."

https://twitter.com/iosif_lazaridis/status/1577030006160515072?s=46&t=55ZrGA4HMJZCZmLzIkUiWQ

After the old Empuries samples and the Iron Age Greek samples of the recently published study, we find again the Mycenaean-like profile in Post-Mycenaean Greeks. It seems that this was the Hellenic profile until the Hellenistic era
 
But Thracians are in the Greek cluster? I wonder what they mean with "Balkans"?

The paper says "[FONT=&quot]Testing a wide range of possible BA and IA sources with [/FONT]qpAdm, valid models of ancestry consist of mixtures of one source related to central or eastern Mediterranean groups (Sicilian, Aegean, or Balkan) and one source related to central or western European groups (France, Spain, Czechia, or Hungary). A genetic origin in the Balkans is suggested by their Y chromosomes, belonging to the E-V13 lineage, which has its highest modern-day frequency in that region"


Very possibly there is some Thracian ancestry here
 
Lazaridis in twiter

"There are many interesting findings, but I will highlight a few:First, many of the 5th c. BC Greek combatants were genetically very similar to Bronze Age Mycenaean Greeks (a thousand years earlier).

Thus, we may conclude that the Ionian and Dorian ethnē -from which the Sicilian Himerans were drawn, according to historical sources- did not represent the migration of a genetically dissimilar population into post-Mycenaean Greece.

Surprisingly, men of diverse genetic background (neither Greek nor Sicanian) were buried in mass graves in the city's West Necropolis: these men were drawn from the east Baltic, the Eurasian steppe, the Balkans, and the Caucasus."

https://twitter.com/iosif_lazaridis/status/1577030006160515072?s=46&t=55ZrGA4HMJZCZmLzIkUiWQ

After the old Empuries samples and the Iron Age Greek samples of the recently published study, we find again the Mycenaean-like profile in Post-Mycenaean Greeks. It seems that this was the Hellenic profile until the Hellenistic era

Himera and Empuries were both founded by Ionians though. It was the south coast of Sicily that had Dorian colonies. Perhaps they have samples from there too that’s why he sounds so assured.
 
The paper says "Testing a wide range of possible BA and IA sources with qpAdm, valid models of ancestry consist of mixtures of one source related to central or eastern Mediterranean groups (Sicilian, Aegean, or Balkan) and one source related to central or western European groups (France, Spain, Czechia, or Hungary). A genetic origin in the Balkans is suggested by their Y chromosomes, belonging to the E-V13 lineage, which has its highest modern-day frequency in that region"


Very possibly there is some Thracian ancestry here

I doubt they would be able to distinguish and I doubt it cluster with Thracians. They would likely think it's a Greek with a slightly northern pull. Not to mention some Thracians are fully Classical Greek-like.

Now, instead of saying Mycenaean-like we can just say Classical Greek-like or mainstream Classical Greek-like.
 
Quick info about the samples

LocalityContextPrimary genetic affinitySampleDateCoverageY haplogroup
Sicily, PolizzelloSicaniMediterraneanI13383-80078,72%G-Y65* / G2a2b2a1a1c1a1a2
Sicily, PolizzelloSicaniMediterraneanI13385-80078,46%G-Z3428* / G2a2b2a1a1c1a1a2
Sicily, PolizzelloSicaniMediterraneanI13391-80076,25%G-Z1903* / G2a2b2a1a1c1a1(xG2a2b2a1a1c1a1a1)
Sicily, PolizzelloSicaniMediterraneanI13394-80075,57%n/a (female)
Sicily, PolizzelloSicaniMediterraneanI13376-80075,35%n/a (female)
Sicily, PolizzelloSicaniMediterraneanI13381-80074,88%n/a (female)
Sicily, PolizzelloSicaniMediterraneanI13392-80074,49%n/a (female)
Sicily, PolizzelloSicaniMediterraneanI13377-80074,27%n/a (female)
Sicily, PolizzelloSicaniMediterraneanI13386-80074,09%G-Z1903* / G2a2b2a1a1c1a1a2
Sicily, PolizzelloSicaniMediterraneanI13378-80074,08%G-Z1903* / G2a2b2a1a1c1a1(xG2a2b2a1a1c1a1a1)
Sicily, PolizzelloSicaniMediterraneanI13380-80073,47%n/a (female)
Sicily, PolizzelloSicaniMediterraneanI13389-80072,72%G-Z3428* / G2a2b2a1a1c1a1a2
Sicily, PolizzelloSicaniMediterraneanI13387-80071,01%n/a (female)
Sicily, PolizzelloSicaniMediterraneanI13390-80070,71%n/a (female)
Sicily, PolizzelloSicaniMediterraneanI13393-80067,87%n/a (female)
Sicily, PolizzelloSicaniMediterraneanI13395-80063,88%n/a (female)
Sicily, PolizzelloSicaniMediterraneanI13384-80057,09%n/a (female)
Sicily, PolizzelloSicaniMediterraneanI13379-80043,30%n/a (female)
Sicily, PolizzelloSicaniMediterraneanI13382-80036,20%G-Z3428* / G2a2b2a1a1c1a1a2
Sicily, Himera409 BCE battleMediterraneanI7224-65573,57%T-S27463* / T1a2b
Sicily, Himera480 BCE battleNortheastern EuropeI10943-64059,11%R-Y35* / R1a1a1b1a2b
Sicily, Himera409 BCE battleMediterraneanI7225-62872,80%R-L2* / R1b1a1b1a1a2b1(xR1b1a1b1a1a2b1a,R1b1a1b1a1a2b1b)
Sicily, Himera409 BCE battleMediterraneanI7223-62872,77%J-FGC45722* / J2a1a1a2b2a(xJ2a1a1a2b2a1)
Sicily, Himera480 BCE battleMediterraneanI7219-62871,87%G-PF3346* / G2a2b2a1a(xG2a2b2a1a1a1,G2a2b2a1a1a2a,G2a2b2a1a1a4 ,G2a2b2a1a1b,G2a2b2a1a1c)
Sicily, Himera480 BCE battleMediterraneanI7218-59067,24%T-S27463* / T1a2b
Sicily, Himera480 BCE battleBalkans and Central EuropeI10950-59062,34%E-CTS6377* / E1b1b1a1b1a(xE1b1b1a1b1a1)
Sicily, Himera480 BCE battleMediterraneanI10945-59062,33%J-Y151557* / J2a1a1a2b(xJ2a1a1a2b2)
Sicily, Himera480 BCE battleMediterraneanI7217-59061,08%L-L595* / L2
Sicily, Himera480 BCE battleMediterraneanI10948-59055,25%R-Y13200* / R1b1a1(xR1b1a1b)
Sicily, Himera480 BCE battleMediterraneanI10952-59054,48%J-Z7706* / J2a1a1b3
Sicily, Himera480 BCE battleBalkans and Central EuropeI10946-59053,87%E-BY6357* / E1b1b1a1b1
Sicily, Himera480 BCE battleCaucasusI10951-59053,42%R-FGC4547* / R1a1(xR1a1a1b1a2)
Sicily, Himera480 BCE battleCaucasusI17872-5902,78%E-V13 / E1b1b1a1b1
Sicily, Himera480 BCE battleBalkans/Western EuropeI17870-5901,85%F / F
Sicily, Himera480 BCE battleEurasian SteppeI10944-58460,16%N* / N
Sicily, Himera480 BCE battleNortheastern EuropeI10949-58249,19%I-L233* / I2a1a(xI2a1a2)
Sicily, Himera480 BCE battleEurasian SteppeI10947-57661,94%R-Z2124* / R1a1a1(xR1a1a1b1a2a,R1a1a1b1a2b)
Sicily, Himera480 BCE battleMediterraneanI7221-57074,47%G-Z42565* / G2a2a1a2(xG2a2a1a2a)
Sicily, HimeraCivilian populationMediterraneanI17432-55066,63%G-PF3346* / G2a2b2a1a
Sicily, HimeraCivilian populationMediterraneanI20168-55012,86%G-L13 / G2a2b2a1a1a1a
Sicily, HimeraCivilian populationMediterraneanI20166-55011,90%L* / L
Sicily, HimeraCivilian populationMediterraneanI20163-5509,50%R-M269* / R1b1a1b
Sicily, HimeraCivilian populationMediterraneanI20160-5503,87%F / F
Sicily, HimeraCivilian populationMediterraneanI20162-5503,47%R-M269* / R1b1a1b
Sicily, HimeraCivilian populationMediterraneanI20161-5501,55%n/a (female)
Sicily, HimeraCivilian populationMediterraneanI20167-5500,99%n/a (female)
Sicily, Baucina, Monte FalconeSicaniMediterraneanI13125-5006,52%G-Z1903* / G2a2b2a1a1c1a1(xG2a2b2a1a1c1a1a1)
Sicily, Baucina, Monte FalconeSicaniMediterraneanI13128-5005,99%R-FT40455* / R1b1a1b1a1a2(xR1b1a1b1a1a2c,R1b1a1b1a1a2f)
Sicily, HimeraCivilian populationMediterraneanI17878-4502,15%G-Z7016 / G2a2b2b1a1a2
Sicily, HimeraCivilian populationMediterraneanI17887-4501,84%n/a (female)
Sicily, HimeraCivilian populationMediterraneanI17881-4501,71%n/a (female)
Sicily, HimeraCivilian populationMediterraneanI17879-4501,55%n/a (female)
Sicily, Himera409 BCE battleMediterraneanI17866-40964,93%J-Z35779 / J2a1a2b2a2b2~
Sicily, Himera409 BCE battleMediterraneanI17884-4094,56%L-L595 / L2
 
PCA

ZxNH7gD.png
 
@Kuivamaa
"Himera was a colony founded by Ionian and Dorian Greeks around 648 BCE (25, 26) (Fig. 1). Himera was also likely inhabited by indigenous Sicilians, Punic people, and Etruscans (25, 27), as interactions between these groups have been well documented at Greek settlements, such as Selinunte (28)."

"
Beyond the intermarriage between locals and Himera’s Greek settlers, it is important to consider the possibility that the composition of the population might also have been influenced by a large-scale influx of Dorians from Agrigento after a political takeover by the Agrigentine tyrant, Theron, in 476 BCE, during which thousands of Himera’s inhabitants were killed (Diod. 11.49); however, the limited sample size available for analysis at this point and the lack of comparative data from Dorian contexts does not allow us to test this hypothesis. "


 

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