Origins of Greek Civilization: Minoans and Mycenaeans

2.5 qpWave/qpAdm
Florian Clemente, J. Víctor Moreno-Mayar, Anna-Sapfo Malaspinas

See Table 3 and Table S3 for results.
Early Bronze Age Aegean (ca. 2,900-2,400 BC: Pta08, Mik15, Kou01, Kou03)
Pertaining to the analysis of ultimate source, when Anatolia Neolithic was one of the
reference populations, the EBA indviduals (Kou01, Kou03, and Mik15 ) were consistent
with a 2-way admixture model of Anatolian Neolithic farmers (∼ 69 − 85%) and Iran
Neolithic/Caucasus HG individuals (∼ 15 − 31%). Although this model could be rejected for Pta08, the data suggest a close relatedness among all EBA individuals. For
instance, the analysis of Pta08 was consistent with a 2-way admixture model of Minoan
Lasithi/Minoan Odigitria/Mik15 (93-99%) and Iran Neolithic/Caucasus HG (1-7%). Together, these results support the idea of EBA individuals in Greece being descendants
of populations related to Anatolian farmers and the Iran Neolithic/Caucasus HG. The
3-way admixture models based on the ultimate sources further support this picture: when
using Anatolia Neolithic as one of the reference populations, the majority of supported
models (i.e., models which could not be rejected) either involved Caucasus HG or Iran
Neolithic as the second major ancestry component, whereas the third component was
more variable and contributed a lesser amount to the Test population. Note that in the
case of Pta08, Neolithic Anatolia was often replaced by an Aegean Bronze Age population,
e.g. Minoan Lasithi or Minoan Odigitria as the major contributing reference population
(often > 90%). This supports the notion that Pta08, which is likely older than Minoan
Lasithi or Minoan Odigitria, is also a Minoan individual. In general, the successful 3-way
admixture models also reflect the high genetic affinity amongst our EBA individuals
in spite of their cultural differences - Minoan (Pta08 ), Cycladic (Kou01, Kou03 ), and
Helladic (Mik15 ), respectively.
When proximate sources were considered, only Kou01 and Kou03 could be successfully
modeled in terms of Anatolian farmers together with one or two additional sources (aside
from the other BA individuals as potential reference populations). Here, in addition to
Iran Neolithic/Caucasus HG populations, Armenian populations were frequently found
among possible admixture models. This is likely due to Armenians sharing a large portion
of the Iran Neolithic/Caucasus HG component, as shown in the ADMIXTURE results.
Taken together, these results are in line with the ADMIXTURE results and MDS plots,
in suggesting that the Early Bronze Age individuals can be distinguished from Anatolian
Neolithic farmers by the presence of an additional Iran Neolithic/Caucasus HG component.
As the locations of our EBA individuals stretch from Crete, to the Cyclades, to the
peninsula of Greece - and as the Iran Neolithic/Caucasus HG component is also observed
in Anatolia in the ADMIXTURE analysis - it appears likely that this component was
wide-spread around the Aegean and parts of Anatolia by 3,000 BC (Lazaridis et al., 2017).

Logkas individuals (ca. 2,000 BC: Log02, Log04)
For the younger Helladic-Logkas-MBA individuals, we could reject all of the 2- and the
majority of the 3-way admixture models based on the ultimate sources, when Anatolian Ne
olithic farmers were one of the sources and neglecting cases where both Logkas individuals
were involved. Successful models generally involved one of the Logkas individuals as Test
population, and the other as one of the sources, with admixture proportions of up to 98.4%,
confirming the close relatedness of these individuals. However, both Helladic-Logkas-MBA
individuals were consistent with a 3-way admixture model of Anatolia_N, Balkans_LBA,
and a third component, e.g., CHG. Furthermore, we could model the Log04 individual
in terms of Anatolia_N (∼ 58%), EHG (∼ 27%), and CHG (∼ 16%). Note that EHG
and CHG are the main components of the Steppe EMBA genome (Barros Damgaard
et al., 2018), indicating that the Aegean MBA individuals may be the result of admixture
between Anatolian farmers/Aegean EBA individuals with Steppe EMBA populations or
Balkan populations that were shown to have received Steppe gene-flow (Mathieson et al.,
2018).
With the proximate sources, further evidence was observed of Steppe admixture in the
Aegean MBA individuals. In the simplest case, we found the Log04 individual to be
consistent with a 2-way admixture model of Anatolia Neolithic (∼ 53%) and Early/Middle
Bronze Age Steppe (∼ 47%) as well as Anatolia Neolithic (∼ 38%) and Middle/Late
Bronze Age Steppe (∼ 62%). Alternatively, 2-way admixture models of EBA individuals
and Steppe/Europe LNBA/Balkan LBA were among the most frequently successful
models for both individuals. This supports the above hypothesis that all Early Bronze
Age individuals were shaped by a geographically widespread Iran/Caucasus influence,
which later encountered populations from the Steppe on the Greek mainland (Haak et al.,
2015; Olalde et al., 2019).
The results from the 3-way admixture model further support the Steppe influence on the
Aegean MBA individuals. Other potential sources of admixture with Anatolia Neolithic
were mainly Bronze Age Armenians, Europeans, and Balkans. According to the admixture
results, Armenians are similar to Iranian/Caucasus populations, whereas Bronze Age
Europeans and Balkans carry the Steppe component.

The diverse genetic origins of a Classical period Greek army

Significance
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.

The classical Mediterranean world was characterized by long-distance interactions mediated not only by trade and colonization (1–4) but also by political and military altercations often precipitated by the need to defend newly established settlements and trade routes that arose due to the expansionist strategies by Greeks, Phoenicians, and others (3, 5). Violent conflict is a common theme of ancient written accounts, and to better understand these ancient societies, it is important to test alternative hypotheses about the role of military activity in their population interactions (6–8).
There is extensive archaeological evidence for links between prehistoric Sicily and the Eastern Mediterranean, as documented by material culture exchange beginning at least in the Early Bronze Age (4, 9–11). This early contact is also reflected in genomic data showing the appearance of an Iranian-related ancestry component from the mid-second millennium BCE onward (12), a type of ancestry that, up to this period of the Mediterranean, had only been detected in regions further to the East (13–15). The establishment of Phoenician trading posts on the western coasts of Sicily, previously entirely inhabited by local Iron Age ethnic groups, began in the ninth century BCE and was followed by a systematic colonization by Greeks in the eighth century BCE, according to archaeological and textual evidence (3, 9, 10, 16–18). Although published ancient DNA data from the Greek world is still sparse, data are available from Late Bronze Age (LBA) people who lived in the Aegean, sometimes referred to as “Mycenaeans”, who are consistent with being the primary source population for present-day people in Greece (13). Inhabitants of the Greek colony of Empúries in Iberia living in the fifth century BCE have clear genetic linkages to LBA Aegean people as well as locals (19), a contribution to people of the island by people with this LBA Aegean genetic signature (20). However, beyond that, little genetic data exist to evaluate diversity of Greek colonies, despite extensive archaeological evidence for diverse identities in Greek settlements throughout the Mediterranean and especially on Sicily (3, 10, 16, 17, 21–24).
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). Himera was the westernmost Greek settlement in northern Sicily (as Selinunte was in the south) and was adjacent to nearby indigenous Sicilian and Punic settlements (18, 29, 30). Ancient authors describe two battles fought at Himera in 480 BCE and 409 BCE between Greeks and Carthaginians, and mass graves containing adult males unearthed in Himera’s West necropolis are believed to contain soldiers who died in the two battles (31–33) (SI Appendix, Figs. S1–S7 and Table S1). According to written sources, Himera won the battle of 480 BCE, owing to relief forces sent by its Greek allies in Syracuse and Agrigento (Diod. 11.20–25) (34), but lost the battle of 409 BCE, when Himerans fought unaided (Diod. 13.60–62; Xen. Hel. 1.1.37) (35). Himera was destroyed and ceased to function as an autonomous city-state (polis) after its loss in 409 BCE.