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Post by Admin on May 4, 2021 21:05:26 GMT
Assessing sex-biased gene flow and inbreeding during the EBA and MBA To assess sex-biased gene flow among the BA Aegeans, mtDNA, Y-, and X-chromosomes were analyzed. The 17 inferred mtDNA (Tables 2 and S2) and the two Y-chromosome haplogroups (Table 2) are common among European Neolithic individuals and do not show any clear evidence of sex-biased gene flow from outside of the Aegean (Document S1). To further investigate sex-biased gene flow, we compared the ancestry on the X chromosome versus the autosomes with a supervised ADMIXTURE following Goldberg et al., 2017. We found no evidence for sex-biased gene flow in EBA Aegeans, with point estimates of Iran_N/CHG-like ancestry on the X chromosome overlapping with those of autosomes (Figure 5A). In contrast, among MBA Aegeans, although Log04 has similar amounts of Steppe-like ancestry on the X chromosome and the autosomes, Log02 is inferred to harbor no Steppe-like ancestry on the X chromosome versus 25%–52% Steppe-like ancestry on the autosomes (Figure 5B). Moreover, in the mtDNA, we found no significant (STAR Methods) population structure (AMOVA p value = 0.293) between EBA and MBA Aegeans from the North of Greece (Pella, Paliambela, Xeropigado Koiladas, and Elati-Logkas) (Figure 1; Document S1). Together, these patterns on the X chromosome and mtDNA could be explained by male-biased gene flow from Steppe-like ancestry into the Aegean. Similarly, Goldberg et al., 2017 and Olalde et al., 2019 suggested that the immigration of Pontic-Caspian Steppe populations during the Late Neolithic/EBA in Europe may have involved a much larger number of males than females.  Figure 5 Sex-biased gene flow Comparison of X-linked and autosomal genetic ancestries associated with (A) Iran_N/CHG-like and (B) Steppe-like components in EBA and MBA Aegean ancient genomes. Violins show the distribution of point estimates across 100 replicates for the corresponding autosomal ancestry, with median indicated by a dot and interquartile ranges indicated by boxes. For the violin plots, we considered a random set of autosomal SNPs matching X-linked SNPs in number (i.e., 8,133) (STAR Methods). Orange diamonds show the point estimates and one associated Standard Error for the same ancestries on the X chromosome. To gain further genetic clues about marital practices during the EBA and the MBA, we inferred contiguous genomic regions in homozygous states—also called runs of homozygosity (ROH)—in four present-day Greek and the six BA Aegean whole genomes (Figure S7). Log04 had more (twenty-nine versus seven at most) and longer ROH (two ROH above 5 Mb) (Document S1) than other ancient individuals. Different evolutionary/demographic processes (Ceballos et al., 2018; Pemberton et al., 2012), including recent inbreeding (Yengo et al., 2019), could explain the Log04 data; in any case, Log02 does not harbor similarly long ROH, suggesting that the underlying cause may not generally characterize the Helladic-Logkas-MBA (Document S1).  Figure S7 Estimated total ROH length by size category for six ancient and four modern genomes, related to Document S1 (A) ROHs estimated from 43 million imputed transitions and transversions, and (B) 13 million imputed transversions (STAR Methods). LBA Myceneans: Armenia versus Steppe-like gene flow The last phase of the BA is associated with a Late Helladic culture termed Mycenaean. Around 1,200 BCE, the Mycenaean civilization began to decline, the palaces were destroyed, the system of writing (Linear B) was abandoned, and their arts and crafts ceased. The causes of their decline are disputed (e.g., climatic change, invasions) (Middleton, 2020). Lazaridis et al., 2017 showed that Mycenaeans were quite distinct from present-day populations, but it remained unclear how they relate to EBA populations. Despite cultural similarity with the Helladic-Logkas-MBA individuals, analyses suggest that the Mycenaean-Peloponnese-LBA were quite distinct genetically, occupying a position in-between the Logkas and the EBA Aegean and the Minoan-Lasithi-MBA in MDS (Figure 2). Unlike the Logkas individuals, they carry a lower European-HG-like component in ADMIXTURE (Figure 3) and do not share significantly more alleles with Iran_N/CHG or EHG compared to Anatolia_N in the D-statistics (Figure S6). However, like the Helladic-Logkas-MBA, they share more alleles with Steppe_EMBA. Mycenaean-Peloponnese-LBA had previously been shown to be consistent with a qpWave/qpAdm model that either involved BA Steppe- or Armenian-related populations (Lazaridis et al., 2017). We recapitulated this result and we additionaly found that Mycenaean-Peloponnese-LBA data are also consisent with a model involving an EBA Aegean and Anatolia_N as source populations (Table 3). In contrast, the Helladic-Logkas-MBA require a Steppe-like source and cannot be explained with a simple model involving an Armenian-like source (Tables 3, S3, and S5). There are further alternative explanations consistent with the data. First, the Mycenaean-Peloponnese-LBA could be the descendants of populations closely related to the MBA Logkas population and to an EBA Aegean population—a 2-way admixture between populations related to Helladic-Logkas-MBA (∼21%–36%) and the Minoan_Odigitria_EMBA and Minoan_Lasithi_MBA (∼64%–79%). Similarly, a 2-way admixture between the Helladic-Logkas-MBA Log04 individual (∼34%–36%) and EBA Aegeans (∼64%–66%) could not be rejected (Table S3). Second, populations related to Armenia BA may have contributed to the Aegeans in a geographically localized fashion during the LBA or earlier (Table S5). This scenario was proposed in the archaeological literature (Drews, 1988) and would imply that the Mycenaeans would not have left much trace in individuals from later generations. Present-day Greek populations resemble MBA Logkas Lazaridis et al., 2017 found present-day Greek populations to be quite distinct from later phases of the BA in the Aegean. In contrast, our results reveal that present-day individuals from Greece (northern Greece—Thessaloniki—and Crete) are closely related to the Helladic-Logkas-MBA individuals of northern Greece, falling near present-day Greeks in MDS analysis (Figure 2), sharing the same ancestry components in ADMIXTURE (Figure 3), and having very similar D-statistics (Figure S6). Moreover, in qpWave/qpAdm analyses (Table 3), the Thessaloniki individuals could be successfully modeled with ∼93%–96% MBA Logkas-related ancestry, and a small fraction (4%–11%) of a second component (either EHG or Eurasian Upper Paleolithic populations such as Kostenki14 [Fu et al., 2016] or MA1 [Raghavan et al., 2014]). The latter are basal populations that constitute a distant outgroup to the Aegean genomes and appear to be interchangeable in this analysis across tests. This suggests that modern populations from northern Greece and Crete could be descendants of Aegean EBA populations, with subsequent admixture with populations related to the Pontic-Caspian Steppe EMBA. Interestingly, modern Cypriots carry no evidence for Steppe-like gene flow across analyses (Figures 2, 3, and S6; Table 3). |
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Post by Admin on May 4, 2021 22:50:54 GMT
  Table S1 Phenotypic insights: Pigmentation and lactose intolerance Using genotype data, we predicted that Pta08, Kou01, and Log02 most likely had brown eyes, dark brown to black hair, and dark skin (Table S1; STAR Methods). These predictions match the visual representations of male individuals from BA wall paintings of Minoan Crete for hair and eye color. The eye and hair color predictions were similar to those from later periods of the Aegean BA (Lazaridis et al., 2017). Although the overall prediction for all three individuals was of dark skin, they also all carried alleles strongly associated with lighter skin color (rs1426654 in the gene SLC24A5, and rs16891982 in SLC45A2) (Mathieson et al., 2015). The latter is in line with observations that skin depigmentation has been segregating since the Neolithic in southern Europe (Hofmanová et al., 2016; Mathieson et al., 2015). Adulthood lactose tolerance was tested on two strongly associated variants, the T allele for rs4988235 under selection in ancient and modern Europeans (Enattah et al., 2008; Mathieson et al., 2015; Tishkoff et al., 2007) and the A allele for rs182549 (−22018A) (Enattah et al., 2002). All three individuals carried the ancestral state in homozygous form (including the MBA Logkas) both at −13910T and at −22018A. This is in line with other results for Neolithic Europeans and Aegeans (Allentoft et al., 2015; Hofmanová et al., 2016; Mathieson et al., 2015) and suggests that dairying may well have been practiced (Evershed et al., 2008) while individuals were lactose intolerant (Document S1). This observation supports a model of mutation-limited adaptation, as has been observed widely across species and phenotypes (Casillas and Barbadilla, 2017; Harris et al., 2018; Jensen et al., 2019). Phenotype prediction The genotype likelihoods from the mapped reads of our nuclear capture data were calculated using ANGSD v. 0.921 (Korneliussen et al., 2014). For this, we specified the SAMtools model (-GL 1) and inferred major/minor alleles from the genotype likelihood (-doMajorMinor 1 -doMaf 1). We used a minimum read depth of 20 (-genoMinDepth 20) and trimmed 10 bases of the reads from both ends (-trim 10). Furthermore, we assumed a uniform prior for the genotypes (-doPost 2) and only considered called genotypes with posterior probabilities > 0.95 (-postCutoff 0.95) and base (-minQ 30) and mapping (-minMapQ 30) qualities of 30 in Phred score. Using these data, we extracted genotypes for Kou01, Log02 and Pta08 for two SNPs on chromosome 2 – rs4988235 and rs182549 – in which the alleles T −13,910 and A −22,018 respectively are informative of lactase persistence. Furthermore, we used the capture data to infer eye-, hair- and skin color for the individuals Kou01, Log02 and Pta08 using the HIrisPLex-S DNA Phenotyping Webtool (https://hirisplex.erasmusmc.nl/). We compared our dataset with the 41 SNPs published as HIrisPlex-S (Chaitanya et al., 2018; Walsh et al., 2017) and found an overlap of 30 SNPs (Table S1, “Nuclear capture” tab). Note that HIrisPLex-S is an extension of the previously published HIrisPLex, which contains 24 SNPs for eye and hair color determination (Walsh et al., 2014) and that our data cover 23 of those 24 SNPs. The HIrisPLex-S webtool calculates individual prediction probabilities and associated values for the loss of prediction accuracy (AUC loss) depending on the available set of SNPs. The results for Kou01, Log02 and Pta08 are given in Table S1. The most supported eye, hair and skin color is the one with the highest prediction probability (Table S1). A discussion about lactose intolerance can be found in Document S1. Concluding remarks During the EBA, the Aegean saw key innovations in trade, craft specialization, social structure, and urbanization. These changes—that mark the end of the Neolithic Period—left indelible marks on Europe and signaled the start of the urban revolution. At the beginning of this cultural transformation, the Aegean world was mostly split between three iconic palatial civilizations, the Helladic, the Cycladic, and the Minoan, each distinguishable by their artwork, pottery style, burial customs, and architecture (Cline, 2012; Shelmerdine, 2008). To better understand the origin of the people behind this transformation, we sequenced four EBA individuals covering all three Aegean BA cultures (Helladic, Cycladic, and Minoan), two MBA individuals from northern Greece, as well as 11 mtDNA genomes from EBA Aegeans. The increased number of variants covered by the whole genomes from this study compared to previous SNP capture data from later periods in the BA Aegean (Figure S2A), as well as the inherent random variant selection characterizing whole genomes (Figure S2B), allowed us to perform demographic inference and statistically contrast population histories. Moreover, the whole genomes generated here can be easily combined with any genomic data (whole genomes, capture data—1240K or otherwise) with a limited loss of variants in future studies of human population history. Note that future work will be required to determine how representative the analyzed genomes of the Aegeans are of the BA Cycladic, Minoan, and Helladic cultures as a whole. In summary, these genomes from the Cycladic, Minoan, and Helladic (Mycenaean) BA civilizations suggest that these culturally different populations were genetically homogeneous across the Aegean and western Anatolia at the beginning of the BA. The EBA genomes drew their ancestry mainly from local Aegean farmers and from populations related to the CHG. These findings are consistent with long-standing archaeological theories regarding the Neolithic-Bronze Age transformation, namely the immigration of new peoples from Anatolia and the Caucasus (Blegen and Haley, 1928; Caskey, 1971; Wace, 1957). However, because the contribution of the local Neolithic populations was significant (Dickinson, 2016; Renfrew, 1972; Tsountas and Manatt, 1897), both local and incoming elements appear to have contributed to the EBA innovations. In contrast, the MBA Aegean population was considerably more structured. One likely reason for such structure is additional Pontic-Caspian Steppe-related gene flow into the Aegean, for which evidence was seen in the newly sequenced MBA Logkas genomes. Present-day Greeks—who also carry Steppe-related ancestry—share ∼90% of their ancestry with MBA northern Aegeans, suggesting continuity between the two time periods. In contrast, LBA Aegeans (Mycenaeans) may carry either diluted Steppe- or Armenian-related ancestry (Lazaridis et al., 2017). This relative discontinuity could be explained by the general decline of the Mycenaean civilization as previously proposed in the archaeological literature (Middleton, 2019). Finally, the inferred migration waves all predate the appearance of Linear B script (1,450 BCE) (Chadwick, 2014). As a result, the genomic data could support both dominant linguistic theories explaining the emergence of Proto-Greek and the evolution of Indo-European languages (Gray et al., 2011). Namely, that these languages either originated in Anatolia (Renfrew, 1972, Renfrew, 1989, Renfrew, 2000) (correlating with the Anatolian and Caucasus-like genetic ancestries) or they originated in the Pontic-Caspian Steppe region (Anthony, 2010) (correlating with the Steppe-like ancestry). Future Mesolithic to BA genomes from Armenia and the Caucasus regions in general could help to further pinpoint the origins and the mode of gene flow into the Aegean and to better integrate the genomic data with the existing archaeological and linguistic evidence. |
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Post by Admin on May 5, 2021 5:53:55 GMT
Document S1. Archaeological background and population genetics.
Related to STAR Methods. Further information about the Final Neolithic and Early
Bronze Age in the Aegean, the importance of the Aegean Bronze Age cultures for Europe,
5 migration theories for the "coming of the Greeks" and major cultural changes in the
Early Bronze Age and Middle Bronze Age, the "coming of the Greeks" based on linguistic
hypotheses, archaeological sites as well as the mitochondrial DNA, the Y chromosome, the
multidimensional scaling analysis, the admixture analysis, the qpWave/qpAdm analysis,
the runs of homozygosity, the ABC-DL analysis and the lactose intolerance analysis
1 Archaeological background
Christina Papageorgopoulou, Elissavet Ganiatsou, Georgia KaramitrouMentessidi, Olga Philaniotou, Adamantios Sampson, Dimitra Theodorou,
Metaxia Tsipopoulou, Ioannis Akamatis, Paul Halstead, Kostas Kotsakis,
15 Dushka Urem-Kotsou, Diamantis Panagiotopoulos, Christina Ziota, Sevasti
Triantaphyllou
1.1 Final Neolithic and Early Bronze Age in the Aegean
The transition from a foraging hunting and gathering subsistence, to a sedentary way of
life, known as the Neolithic Revolution, was a process that decisively changed human
history (Bellwood, 2005). The primary innovations of the Neolithic Revolution included
permanent settlements, as well as plant and animal domestication. It has been well
established that these innovations spread from southwestern Asia and reached Europe
at the beginning of the 7th millennium BCE (Renfrew and Bahn, 2014). This spread
was mediated by the expansion of farmers into Central and Western Europe (Hofmanová
et al., 2016; Mathieson et al., 2018). During this expansion, the Aegean region played an
important role in providing a stepping stone between Anatolia, Central Europe and the
Western Mediterranean (Theocharis, 1973; Özdoğan, 2006; Kotsakis, 2008).
The first Neolithic communities in Greece appeared between 6,700 and 6,600 cal BCE,
as revealed by archaeological data (Douka et al., 2017; Karamitrou-Mentessidi et al., 2015;
Maniatis, 2012; Perlès et al., 2013) (Tab. i). The economy of these early communities
appears to have been exclusively based on farming and herding. Pottery, clay stamps,
and schematic figurines were present from the beginning of the Neolithic period (Perlès,
2001). From the Middle Neolithic (5,600-5,300 BCE) onward, a remarkable increase in the
number of settlements, even in less favorable environments, has been recorded (Demoule
and Perlès, 1993; Andreou et al., 1996). The formation of more and larger communities
and the use of secondary, renewable products from animals and plants (such as traction
and milk - known as secondary products (Sherratt, 1981)), promoted more complex social
structures and larger economic networks, as appeared during the Late Neolithic period
(5,300-4,500 BCE).
These phenomena became more pronounced during the Final Neolithic (4,500-3,200
BCE), also called Chalcolithic because of the first use of metals1 . During this period a transition from farming and herding, to the more advanced economy of the Early Bronze Age (EBA), was unfolding. Production activities improved, maritime contacts and trade exchange were intensified, and mobility for the provision of metals had begun. The coastal
zones and the islands were now populated and in the mainland certain settlements seem
to have acquired considerable economic importance (Broodbank, 2008).
However, during the Final Neolithic period, throughout the Balkan peninsula an
abrupt termination of habitation has been observed (for a different view see Kotsakis,
2018; Renfrew, 2018). This is particularly evident in southeastern Bulgaria and southern
Romania, where tell sites like Karanovo, Yunatsite, Goljamo, Delcevo, Ovcarovo, Hotnica
are abandoned (Todorova, 1995). Similarly, in Greece, the archaeological record reveals
a discontinuity for several centuries (500–1,000 years), in tell sites like Servia, Mandalo,
Sitagroi, Dikili Tash, flat settlements (Makriyalos, Vasilika) and caves (Alepotrypa,
Diros) (Manning, 1995; Andreou, 2012; Tsirtsoni, 2016b). Diverse causes for this abrupt
termination of habitation have been suggested, including 1) climatic changes (Lespez
et al., 2016; Todorova, 1995; Weninger et al., 2009), 2) a breakdown of traditional social
structures in communities (Chapman et al., 2006; Windler et al., 2013), and/or 3) invasions
of Proto-Indo-European-language groups from the Pontic-Caspian Steppe (Gimbutas,
1956; Coleman, 2000) also debated by other researchers (Andreou, 2012; Whittle, 1997).
Paleogenomic studies have recently revealed the influence of such groups on Central and
Southeast Europe (Allentoft et al., 2015; Mathieson et al., 2015; Mathieson et al., 2018).
This disruption of habitation was followed by cultural changes, which are more evident in
northern Greece (e.g., Macedonia). In the south, the material culture remained unchanged
between the last phases of the Neolithic and the beginning of the Early Bronze Age (EBA)
(Tsirtsoni, 2016a; Pullen, 2008). This continuity was more pronounced in the Cyclades
and Crete, and in areas that had been settled late in the local Neolithic sequence (e.g.
the Northeastern Aegean islands) (Tsirtsoni, 2016a).
The Final Neolithic, in general, provided the key features for the development of the
next major cultural transformation, that of the Bronze Age (BA). The BA in Greece
lasted from roughly 3,200 to 1,100 BCE, and it is divided into three main chronological
groups: Early, Middle, and Late BA (Tab. i). The EBA spanned from 3,200 to 2,000
BCE and is characterised by the appearance of early forms of urbanization (Childe, 1942;
Childe, 1950; Renfrew, 1972), extensive exchange networks known as the International
Spirit of the Aegean, and fine craftsmanship. Renfrew (1972), in his influential book "The
Emergence of Civilization in the Aegean" described how the appearance of agricultural
specializations (e.g., wine and oil) led not only to simple production, but also to craft
specialization, redistribution systems, trade, and population growth.
During the EBA the communication networks between the Aegean islands, Crete, and
the coastal regions of Greece and Western Anatolia were extensive. The presence in Crete
of stone artefacts, figurines, and ceramic from the Cyclades and Peloponnese signifies the
intensive trade as well as the important influence of the Cycladic culture on pre-palatial
Crete (Tomkins and Schoep, 2012). The extensive exchange networks have been also
documented in the Aegean, through a wide distribution of artefacts, including the Kampos
group pottery, the folded arm figurines, the mid-rib metal daggers, and the long types of
obsidian blades (Broodbank, 2002). The mortuary treatment (concerning both associated
artefacts and burial practices) indicate a strong influence of the Cyclades over the Greek
mainland and Crete. This is observed in the cemetery of Agios Kosmas in Attica, the
cemetery of Manika in Euboea, the Tholos tomb Γ in Archanes, the cemeteries in Pseira
and Gournes, the cemetery of Hagia Photia and to a less degree the one at Kephala
Petras, in Siteia in north-eastern Crete. The early signs of urbanization accompanied by
large fortified settlements and significant craft specialization appears in the middle of the
Aegean (Strofilas on the island of Andros; Palamari on the island of Skiros; Palioskala in
Thessaly) (Televantou, 2008). Palamari on the island of Skyros show signs of urbanization
(Parlama, 1992) very similar to those of Liman Tepe (Izmir, Western Anatolia), but also
close relationships with mainland Greece and the cities of Poliochni on the island of
Limnos and Troy (Kouka, 2002).
Several settlements developed into strong centers of trade and craftsmanship. The
Aegean provided a fruitful ground for the emergence of the significant BA civilizations,
known as the Aegean Cultures (Treuil, 1989; Dickinson, 1994; Shelmerdine, 2008; Cline,
2012). This term is used to characterize the civilizations of the four geographical areas
that surround the Aegean sea: Minoan Civilization of Crete, Helladic civilization
of mainland Greece, Cycladic civilization referring to the Cycladic islands, and the
western Anatolian cultures (Cline, 2012). These civilizations emerged, evolved and
acted almost in parallel (Tab. i), for a detailed chronology and terminology on Aegean
cultures see Manning, 2012).
During the first period of EBA, economic centers established a surplus of agricultural
products and raw materials, large storage facilities, and redistribution systems. This
triggered social stratification and allowed certain groups of people to exercise leadership
(Gilman et al., 1981). These transformations steadily led to the socioeconomic systems
of the late Middle Bronze Age (MBA) and the eminent Palatial Period. It is during
the MBA Period that the palatial civilizations of Crete and Mycenae reached their peak
with elaborate architecture, fine art, and craftsmanship. During this period, the first
written languages also appear in Europe, the undeciphered Linear A used by the Minoans
(1,850-1,400 BCE) and the deciphered Linear B used by the Mycenaeans (1,400-1,200
BCE) (see Section 1.4). The latter is the earliest attested form of the Greek language
(Ventris and Chadwick, 1953).
Table i: Timeline of Neolithic and Bronze Age cultures in the Aegean
Neolithic period 6,700-3,000 BCE
Early Neolithic 6,700-5,600 BCE
Middle Neolithic 5,600-5,300 BCE
Late Neolithic 5,300-4,500 BCE
Final Neolithic 4,500-3,200/3,000 BCE
Bronze Age Period 3,200/3,000-1,100 BCE
Early Bronze Age 3,200/3,000-2,000 BCE
Early- Helladic/Cycladic/Minoan I 3,200/3,000-2,600 BCE
Early- Helladic/Cycladic/Minoan II 2,600-2,300 BCE
Early- Helladic/Cycladic/Minoan III 2,300-2,100/2,000 BCE
Middle Bronze Age 2,100/2,000-1,600 BCE
Middle- Helladic/Minoan I 2,100/2,000-1,800 BCE
Middle- Helladic/Minoan II 1,800-1,700 BCE
Middle- Helladic/Minoan III 1,700-1,600 BCE
Late Bronze Age 1,600-1,100 BCE
Late Helladic/Minoan I 1,600-1,500/1,450 BCE
Late Helladic/Minoan II 1,500/1,450-1,400 BCE
Late Helladic/Minoan III 1,400-1,100 BCE
Sub-Mycenaean/sub-Minoan: 1,100/1,050-1,025 BCE
source: Hellenic Ministry of Culture
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Post by Admin on May 5, 2021 19:55:57 GMT
1.2 The importance of the Aegean Bronze Age cultures for Europe
The EBA signifies the introduction of new raw materials and techniques (e.g., use of
metals), but also social, political and economic changes. These new social and economic
forces enabled the development of a new system of production-distribution-consumption
and thereby set the roots of modern capitalism (Kristiansen, 2016). The demand for metals
developed an unprecedented need for raw materials, while the invention of technologies,
such as the sail and the wheeled vehicles, facilitated the transportation of the raw materials
and their products.
Systematic trade in tin, copper, textiles and salt had the same significance in the
Bronze Age as gas and oil in modern economies (Kristiansen, 2016). The exchange of such
commodities required networks of producers, traders and consumers, but also distant political alliances, emporia (small cities serving as trading centres) and hierarchical institutions
of governance. Archaeological evidence identifies the Near East and Aegean, as the regions
where the complex societies, that fueled the systematic production, consumption and
redistribution of staple and luxury materials, first developed. Besides its immediate local
significance, this process enabled west and north European communities to interact with a
global economic world, and although they did not immediately adopt such BA innovations
135 as urbanization and writing, they acquired a novel value system still prominent today in
long-distance political treaties and alliances, merchantile economy and regional identities
(Kristiansen, 2016).
1.3 Migration theories for the "coming of the Greeks" and major
cultural changes in the Early Bronze Age and Middle Bronze Age
Based on extensive archaeological data, several hypotheses have been proposed for the
origin and development of BA Aegean cultures, including: 1) local innovation, where
changes were based on genetic and cultural continuity of local Neolithic groups (Tsountas
and Manatt, 1897; Dickinson, 2016; Renfrew, 1972); 2) the immigration of new populations
145 from Anatolia and the Caucasus during the EBA and MBA (Blegen and Haley, 1928;
Caskey, 1971; Wace and Blegen, 1916); and 3) the arrival of probable speakers of IndoEuropean languages
from the Pontic-Caspian Steppe at the beginning of the EBA (for a review, see Coleman, 2000; Pullen, 2008; Dickinson, 2016).
Cultural changes in the Aegean and the appearance of the proto-Greek language (by
1,400 BCE as seen in the Mycenaean Linear B script) have fuelled the debate on migration
and the so-called "coming of the Greeks" (Tab. ii). This debate has concerned European
archaeology for over 120 years and has been intimately linked with conjectured migrations
into Europe from the Pontic-Caspian Steppe region at the end of the Neolithic Period
and their connection to the spread of the Indo-European languages. BA Greece plays a
155 significant role in the Indo-European problem since Greek is one of the oldest attested
Indo-European languages (Gray et al., 2011).
From an archaeological perspective (for a linguistic perspective see Section 1.4), precise
cultural changes and cultural gaps have been linked to the Indo-European hypothesis
and to the appearance of the Greek language and the BA civilizations (Demoule, 2017;
160 Giannopoulos, 2012). This section presents, in summary, the most significant cultural
changes that have triggered debates on the probable arrival of population groups into the
Aegean during the BA. This summary may be juxtaposed to the genetic signals produced
by the present study.
It must be borne in mind, however, that 1) cultural changes are not necessarily the
product of migration; and 2) migrations, are not always identifiable archaeologically2
Moreover, the scenarios presented have been heavily influenced by the theoretical dialogue
between migrationism (cultural change explained by migration) and diffusionism (cultural
change is explained by diffusion of ideas and materials) that has prevailed in archaeology
for the last century.
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Post by Admin on May 5, 2021 20:38:27 GMT
1.3.1 Early Bronze Age
A. Late Neolithic - Early Bronze Age scenario
The first chronological time-point that has been linked to the "coming of the Greeks" is
the last phase of the Neolithic Period (Final Neolithic or Chalcolithic) and the beginning
of the EBA. As described in the previous section the end of the Neolithic in some parts of
(especially northern) Greece, is characterized by large-scale depopulation and abandonment
of established sites. Coleman, after studying the archaeological and linguistic evidence,
suggested that shortly before the beginning of EBA (3,800-3,200 BCE) people from
the North Pontic Steppe arrived in this sparsely populated Greek mainland (not in the
Cyclades and Crete). The newcomers in conjunction with the local cultures founded
the Early Helladic (EH) culture of mainland Greece (Coleman, 2000). He also accepts
Gimbutas’s theory (Gimbutas, 1997), in setting this arrival in waves, the first arriving at
about 4,400 BCE and the second around 3,800-3,200 BCE.
Coleman bases his argument for migration from the North (not further specified) at
the end of the Final Neolithic and the beginning of EBA,on the following points: 1) the
abandonment of sites during the Final Neolithic, the appearance of a small number of EH
I sites and the significant increase in EH II sites (see Tab. i), suggesting that the initial
settlers were successful and the population increased; 2) changes in pottery, mostly in
shapes (for a different view see Sampson (1981) and Demoule (2017)); 3) changes in lithic
technology; 4) increased exploitation of metals; 5) rare use of figurines compared to the
Final Neolithic period (Coleman, 2000; Coleman and Facorellis, 2018).
Arguments 1 and 2 are widely accepted by academic scholars (for counter-arguments
see Bintliff and Sarri (2018) and Kotsakis (2018)) although the interpretation of the
evidence differs significantly. For the apparent depopulation of Greece during the end
of the Neolithic period there is still no persuasive explanation (see section 1 above).
Epidemics, climatic changes and invasions have been suggested. The epidemic scenario
has not been strongly supported, although palaeogenomic evidence for the presence of
plague (Yersinia pestis) already during the Neolithic (Rascovan et al., 2019; Rasmussen
et al., 2015) may be confirmed by future studies. Climatic changes are not verified by
palaeoclimatologists and invasions are largely disputed.
Other scholars have characterized pottery styles as stable, with no significant change
from the Late Neolithic to EH I and no significant rupture in material culture that need
to be explained by the arrival of foreign groups (Demoule, 2017). They underline the
view of Mallory, who sets the geographical limit for the Steppe expansion in Hungary
and finds little support on the view that Steppe invaders arrived in Greece and Anatolia
(Mallory and Adams, 1997). They acknowledge, however, that a common pottery shape
and decorated motif appeared around the 4th millennium BCE in most parts of the Balkan
peninsula. This common style extended from central Europe (Baden culture) and southern
of Hungary, Bulgaria (Ezero culture) to Greece and western Anatolia (e.g., Kumtepe)
(Demoule, 2017; Maran, 1998). Coleman (2000) links this common stylistic relationship
with the arrival of people from the Pontic-Caspian Steppes, largely following previous
theories (Gimbutas, 1997).
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