Origins of Greek Civilization: Minoans and Mycenaeans

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Origins of Greek Civilization: Minoans and Mycenaeans Oct 20, 2022 5:58:57 GMT

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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).
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.

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Origins of Greek Civilization: Minoans and Mycenaeans Oct 21, 2022 7:21:46 GMT

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Integration of Isotopic Evidence Supports Mercenary Presence in the 480 BCE Battle.
While the genetic makeup of a person gives insight into the origin of his or her ancestors, whether recently or many generations back in time, it does not provide information about the geographic region where the person lived or grew up. Strontium and oxygen isotope ratios reflect the local geology and hydrology of a person’s place of residence during the period of tooth or bone formation and provide information about individual mobility in a single lifespan (73). Thus, genetic and isotopic information provide complementary information about human mobility and life history.
We correlated the diversity in genetic histories with strontium and oxygen isotope data gathered from the teeth of Himeran individuals in previous research (37). For all the individuals from the civilian sample, all the individuals from 409 BCE graves, and most of the individuals in the 480 BCE graves determined to be Aegean related (five of seven), the strontium isotopic ratios are consistent with local origins (37). In contrast, all 480 BCE soldiers with primary genetic affinities with central Europe, northeastern Europe, the Eurasian steppe, and Armenia are isotopically nonlocal (Fig. 3). The low oxygen ratios of some individuals are characteristic of higher latitudes, higher altitudes, cooler climates, greater rainfall, and greater distances from coasts than Himera, and the high strontium isotope ratios of some individuals indicate more ancient underlying rocks and soils than at Himera (37, 74). The outlying isotopic data by themselves do not prove non-Sicilian origins for the individuals from Himera: similar outlying ratios for both isotopes can also be found in other parts of Sicily, albeit not in the immediate region of Himera. However, the agreement between genetic and isotopic evidence, showing that all nine individuals with outlying genetic profiles also have nonlocal isotopic profiles, provide compelling evidence that many soldiers who fought in the battle of 480 BCE came from distant regions of the known Greek world.

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Origins of Greek Civilization: Minoans and Mycenaeans Oct 22, 2022 7:00:43 GMT

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Post by Admin on Oct 22, 2022 7:00:43 GMT

Fig. 3.

Summary of isotopic ratios, grave context, genetic affinities, and phenotypic information. Strontium and oxygen isotope data of the soldiers and civilians discussed in this paper. Symbols and symbol colors indicate grave context and genetic affinity, respectively. Associated colored squares indicate from left to right most probable hair, eye, and skin pigmentation (NA = not assessable). Shaded regions represent the local baselines of isotope values for Himera, measured from archaeological fauna in the case of strontium, and from n = 93 humans from Himera’s civilian population in the case of oxygen; see ref. (37) for details on geochemical data.
These findings suggests the possibility that, unlike the genetically homogeneous sampled individuals who died in battle in 409 BCE, soldiers from 480 BCE had more diverse origins. This supports the interpretation that the Greek army in 480 BCE included mercenaries, or paid soldiers, that were of Greek and other cultural backgrounds (38). Both Greek colonies and other polities hired mercenaries to support citizen-based hoplite armies (6, 7, 75–77) (Thuc. 1.60: Greek use of Mercenaries; Hdt. 7.165: Diverse army led by Hamilcar the Carthaginian; Hdt. 8.26: Helping Persia; Hdt.2.152.4–5: Helping Egypt). It is well documented that Punic armies, such as Carthage, often used mercenary armies (8, 42), and it is likely that Greek tyrants on Sicily did the same (43). Although the ancient historians do not specifically mention mercenaries fighting for Himera in 480 BCE, they do mention that the Syracusan tyrant Gelon had gathered a large number of foreign mercenaries to round out his army (Diod. 11.72); these would plausibly have been among the relief forces sent to defend Himera. Sicily was a particularly attractive destination for mercenaries, as many Greek poleis on Sicily were governed by tyrants capable of extracting the kind of wealth needed to fund armies of mercenaries and usually in need of bodyguards to maintain their control (43, 77, 78). Some mercenaries reputedly were recruited from areas famed for particular skills, such as archers from Scythia (36) and peltasts from Thrace (79).
Soldiers from 480 BCE, whose genetic diversity exceeds that of the civilian sample at Himera, therefore likely did not reflect a draft exclusively from the Himeran polis, and the combination of genetic and isotope data suggest they were born and raised even further afield than Sicily itself. In contrast, the isotopic signatures that are mostly consistent with a local origin (37), and the genetic similarity of adult male individuals interred in mass graves associated with the battle of 409 BCE, corroborates historical accounts and suggests a fighting force drawn from Himera’s civilian population, without aid from mercenaries or allies. Historical accounts state that Syracusan Greeks initially were sent to aid Himera in 409 BCE but ultimately turned back in order to protect Syracuse, leaving Himera to fend for itself, relying on its own citizens (Diod. 13.59–61). However, as we only have genetic data for five of the soldiers of the 409 BCE battle, additional data would be required to confirm whether the lack of individuals with non-Aegean genetic origins is statistically significant.

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Origins of Greek Civilization: Minoans and Mycenaeans Oct 23, 2022 4:38:00 GMT

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Post by Admin on Oct 23, 2022 4:38:00 GMT

Identity Markers and Burial Treatment.
There are patterns in genetic affinities among the seven mass graves associated with the battle of 480 BCE. All the soldiers who fall outside the Aegean genetic cluster are interred in mass graves Nos. 1–4 (SI Appendix, Figs. S1–S3), whereas all individuals from mass graves Nos. 5–7 (SI Appendix, Figs. S4–S6) fall within this cluster, a statistically significant difference (P = 0.0028 by a χ2 test with one degree of freedom; Fig. 3). This result mirrors strontium isotope evidence showing more nonlocals interred in mass graves Nos. 1–4 (P = 0.0035 by a χ2 test with one degree of freedom; Fig. 3 and Dataset S2) (37). Mass graves Nos. 1–4 and Nos. 5–7 also are spatially segregated and differ in size, with mass graves Nos. 1–4 comprising significantly more interments (80) (SI Appendix, Figs. S1–S3). The correlation of burial location of individuals with their genetic ancestries suggests deliberate placement of soldiers with similar ancestral backgrounds in the same mass grave, reflecting the importance of identity in determining burial in this society (40, 59, 81).
These results suggest that individuals in mass graves Nos. 5–7 may have had greater prestige among survivors who buried them, consistent with previous arguments based on mortuary context (80). Individuals interred in mass graves Nos. 5–7 are significantly older than individuals interred in mass graves Nos. 1–4 (mean age, 45.0 versus 29.6 y old) and show no signs of having been dragged, as is the case for mass graves Nos. 1–4, or buried hastily, as is the case for mass graves Nos. 8/9 (80). Furthermore, individuals in mass graves Nos. 5 and 6 include grave goods, unlike the other mass graves from 480 BCE (32). The fact that these individuals also fall within the Aegean genetic cluster suggests a link between Aegean ancestry and prestige, as perceived by the individuals responsible for burying the fallen soldiers.
Constructions of ethnicity in the ancient Greek world were chiefly a matter of common descent and ancestral homelands but also were informed by cultural traits, such as language or material culture (82, 83). Our data show that the survivors who interred the casualties of the battle of 480 BCE incorporated information on the social origins of the deceased as part of their social logic for respectful treatment of the dead. Individuals with “foreign” ancestry, all of whom also are identified as nonlocal on the basis of isotopic evidence (Fig. 3 and Dataset S2), were interred in larger mass graves Nos. 1–4, and individuals with genetic affinities to other Greek populations were interred in the smaller mass graves Nos. 5–7. Nevertheless, it is not clear that nonlocals were considered as a homogeneous group of “others”. It is possible that the more-particular origins of the deceased were acknowledged and acted upon; for example, mass graves Nos. 1/2 contain both of the individuals with northeastern European ancestry. Identification of the fallen might have been done by surviving fellow soldiers who knew them, on the basis of their armor and weapons, or on the basis of phenotypic characteristics (Fig. 3 and SI Appendix, Table S19 and Dataset S7). Many other soldiers may have been buried in single interments, rendering them archaeologically invisible as soldiers. At Himera, some single interments do include individuals with perimortem trauma suggestive of being subjects of violence at the time of their deaths. These individuals could have died in battle and been interred in single burials by those who knew them. It is possible that some bodies were repatriated, as is documented in classical antiquity (84–86), with a greater likelihood of repatriation to locations elsewhere in Sicily than to more distant regions. Repatriation would have been achieved by first cremating the body (85, 86), and cremation was commonly practiced at Himera (87).

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Origins of Greek Civilization: Minoans and Mycenaeans Oct 24, 2022 7:02:24 GMT

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Post by Admin on Oct 24, 2022 7:02:24 GMT

Discussion
These results highlight how colonies facilitated migration, as reflected by heterogeneous ancestry deriving from geographically disparate locations. Furthermore, the results show how armed conflict served as a contact mechanism among diverse ancient populations in classical antiquity. In an already highly connected region, Greek warfare facilitated human mobility on a continental scale, providing a new line of evidence that enriches and supports scholarship emphasizing that migration occurred on a variety of scales, including small-scale movements of people (e.g., Horden and Purcell (88)), large-scale movements of groups (e.g., colonists; van Dommelen (89)), and globalization as an ancient process (90). Much of the migration related to warfare in the ancient Greek world may have been routine, small-scale movements of citizen soldiers (i.e., hoplites) (36), as had been documented in other contexts (91). However, our findings support suggestions that warfare, and the involvement of mercenaries, may have played an important role in large-scale movements of individuals in the ancient Greek world (7, 17). Greek mercenary warfare has been a focus of archaeological and historical research (6, 77, 92) but has rarely been discussed as a vehicle for culture change and gene flow in the context of Greek colonies (38), despite the recognized importance of warfare and the involvement of mercenaries as an ethnic unifying force in Phoenician contexts (8, 42, 93, 94). Like merchants, colonists, and slaves, mercenaries played a role in the transmission of culture in the Mediterranean region. While trade and colonization drew together communities around the Mediterranean region in person-to-person interactions (95), exchange of goods across vast geographic space did not necessitate migration and instead could be accomplished through intermediaries (i.e., merchants). Mercenaries were among the most long-distance travelers of the Greek world (rivaled, perhaps, by slaves), bringing persons of widely divergent cultural and genetic backgrounds face to face. Coupling genetic and isotopic evidence for mobility disambiguates the time scale of migration, indicating first generation migrants to Sicily. Mercenaries drawn to the Mediterranean region, and Sicily in particular, from distant regions of Europe, such as the Balkans, Gaul, and Germania, by the economic opportunity to serve Greek tyrants and participate in Greek wars were not merely itinerant “outsiders” in Greek poleis (96). They often lived within local communities, which deeply embedded them in alien cultures, as was the case for merchants and colonists, and rendered them powerful potential actors in cultural exchanges (38, 97). Historical evidence indicates instances of mercenaries becoming enfranchised as citizens and becoming integrated into the political, cultural, and genetic fabric of Greek poleis (75, 78). Diodorus Siculus writes of 7,000 foreign mercenaries in Syracusan tyrant Gelon’s employ (some of whom may have participated in the battle of 480 BCE) who were granted citizenship in 466 BCE (Diod. 11.72.3) (75). Besides Greek settlers and the indigenous peoples they encountered, mercenaries were part of the exchange of culture, ideas, and possibly genes taking place in Greek poleis in the first millennium BCE (38, 97) and might have, in some cases, played an important role in securing Greek military victories, as the divergent outcomes of Himera’s two battles demonstrate.
Historically contextualized genetic and isotopic evidence of foreign soldiers at Himera is another indicator of the heterogeneity and evolving complexity characterizing IA and post-IA societies of Europe, the Middle East, and other world regions (98). Identifying diverse origins of Classical Greek mercenaries provides nuance to our understanding of the long-distance movements of individuals relating to specialized economic or religious activities, apart from well-documented population movements, such as the establishment of colonies. Sites of the ancient Mediterranean for which genetic data have been published vary considerably in terms of settlement type, function, and historical context. The mass graves at Himera are singular in representing armed conflict and providing evidence for warfare as a dispersal mechanism of individuals across vast geographic space. By continuing to evaluate genetic trends as well as unique cases, in archaeological and historical context, research will be able to unravel series of different biographies of localities that, when combined, have the potential to enrich our understanding of trends, such as the intensification of trade, warfare, urbanism, and collapse.