Homogeneity among Illyrian and Greek populations and shared ancestry, study finds

In this post we present the very interesting paper titled “Evidence for Long-Term Migration on the Balkan Peninsula Using Dental and Cranial Nonmetric Data:Early Interaction Between Corinth (Greece) and its Colony at Apollonia (Albania)“, by Britney Kyle McIlvaine et al. We also comment on the findings and their possible further importance.


Abstract This article seeks to identify “Greeks” and “non-Greeks” in “mixed” mortuary contexts in a Greek colony. Specifically, we test the hypothesis that Illyrian and Greek individuals lived and were buried together at the Corinthian colony of Apollonia, Albania (established ca. 600 BC). The pattern of human biological interaction at Apollonia is tested by identifying variation in genetic relatedness using biodistance analysis of dental and cranial nonmetric traits for three sites: Apollonia (n5116), its founder-city Corinth (n569), and Lofkënd (n5108), an inland site near Apollonia pre-dating colonization. Logistic regression analysis estimates that individuals from colonial Apollonia are most closely related to prehistoric Illyrian populations (from Lofkënd and pre-historic Apollonia), rather than Greeks (from Corinth). The phenotypic similarity between colonial Apollonia and prehistoric Illyria suggests that there was a large Illyrian contribution to the gene pool at the colony of Apollonia. However, some trait combinations show low biological distances among all groups, suggesting homogeneity among Illyrian and Greek populations (assessed through pseudo-Mahalanobis’ ). The degree of phenotypic similarity suggests shared ancestry and long-term migration throughout these regions. The impacts of missing data and small sample sizes are also considered.


Despite extensive research on the Greek colonial diaspora (Gwynn, 1918; Graham, 1983; Gosden, 2004; Hodos, 2006; Tsetskhladze, 2006), the extent of inter-breeding between Greek colonists and native peoples remains largely unexplored (van Dommelen, 2012). Lack of knowledge limits our current understanding of the ways that Greek colonists integrated themselves with local populations following colonization. Economic interaction connected spatially and culturally diverse populations throughout the Mediterranean beginning in the Bronze Age, if not earlier (Galaty, 2002; Lafe and Galaty, 2009; Galaty et al., 2010; van Dommelen, 2012). Greek males who participated in trade likely married and interbred with local women (Hodos, 1999; Antonaccio, 2001, 2003; Tsetskhladze, 2006). However, as noted by van Dommelen (2012) the presence of traded “raw materials and finished products, no matter how frequent and long-distance [exchange] may have been, does not by itself imply migration or colonization”. Thus the degree of migration and genetic interaction between Greeks and natives cannot be tested through the material record alone. Using bioarchaeological methods, we test whether the Greeks and local Illyrian populations interbred and were buried together at the Greek colony of Apollonia, Albania.

Unlike the assessment of material culture, which is always indirect, the application of biodistance analysis provides a direct test of whether Greeks, local Illyrians, or a combination of the two groups are buried in the Apollonian cemetery. The current study tests the hypothesis that both Illyrian and Greek individuals were buried together in tumuli (mounds) at the colony of Apollonia, Albania. The hypothesis is tested via biodistance analysis (using phenotypic variation to assess genetic relationships between individuals or populations) of human dentitions and crania. Our approach aims to determine whether skeletons represent native peoples or colonists, and to provide information regarding the extent of migration, followed by non-local interment, for the Greeks.


Corinthian colonization events The first wave of Corinthian colonization in the Mediterranean began in approximately 733 BC with the establishment of Syracuse on Sicily and Corcyra on Corfu (Graham, 1983; Wilkes, 1992; Cabanes, 2008). The Corinthians then expanded into Illyrian territory. The Illyrians, an ethnic group who inhabited large portions of the Balkan Peninsula, are thought to be the ancestors of modern Albanians (Wilkes, 1992; Galaty, 2002). Corinth and Corcyra cofounded the colony of Epidamnus (627 BC) in Illyria (modern Albania) (Tsetskhladze,2006; Cabanes, 2008). Albanian Apollonia may have been established by both Corcyra and Corinth, or by Corinth alone (ca. 600 BC) (Stocker and Davis, 2006;Tsetskhladze, 2006; Cabanes, 2008; Amore, 2010). Archaeologists and historians often describe relationships between Corinthians and Illyrians as mutually beneficial (Hammond, 1992; Wilkes, 1992; Cabanes, 2008; although see Pollo and Puto, 1981), and archaeological evidence, in the form of Illyrian names on many coins and gravestones, suggests a strong and sustained Illyrian presence at Apollonia (Stipcevic, 1977; Galaty, 2002).

At the time Apollonia was colonized, Greeks and Illyrians had distinct burial practices, with Greeks buried in flat cemeteries containing individual inhumations (Galaty, 2002) and Illyrians buried in mounds (i.e., tumuli) with individuals in each tumulus presumably belonged to the same linage or clan (Galaty, 2002). However (as documented in other Greek colonial contexts (Shepherd, 1993, 2005; Leighton, 1999)), a mix of Greek and local burial practices is observed at Apollonia (Amore, 2005, 2010). Following colonization, traditional Illyrian tumuli at Apollonia begin to include large numbers of Greek ceramics and grave goods as well as Corinthian-style limestone sarcophagi (Galaty, 2002; Stocker and Davis, 2006; Amore, 2010). The apparent hybridization of burial traditions contrasts with the burial practices observed at the nearby colony of Epidamnus, where Greek-style flat cemeteries replaced Illyrian burial mounds following colonization (Ceka, 2005). The mix of Illyrian-style tumuli and Greek-style grave goods at Apollonia led archaeologists to debate the ethnic affiliations of individuals in the burial mounds (as discussed in Lafe, 2003; Amore, 2005, 2010). Some propose that the mounds contain exclusively native Illyrians or exclusively Corinthians (Hammond, 1992; Wilkes, 1992; Ceka, 2005). Others suggest the Apollonian burial mounds contain remains of both Corinthian colonizers and local Illyrians (Lafe, 2003; Amore, 2005, 2010; Schepartz, 2010), as originally proposed by Albanian archaeologist Aleksandra Mano (Amore, 2005, 2010).

Materials Skeletal materials from three archaeological sites were analyzed here: the colony of Apollonia, its founder-city Corinth, and Lofkënd, an inland site near Apollonia predating colonization.


Apollonia, Albania Apollonia, located in southwestern Albania approximately 10 km from the coast of the Adriatic Sea (Amore, 2010), emerged as an important trade site following colonization in the beginning of the 6th century BC (Stipcevic, 1977; Wilkes, 1992; Amore, 2005, 2010; Stocker and Davis, 2006), when 200 Corinthians “established good relations with the local Illyrians [and] founded a joint settlement” (Hammond, 1992; p 31-32;Cabanes, 2008). The Apollonian skeletal series was excavated between 2002 and 2006 by M.G. Amore (2010), and burials were recovered from the prehistoric Bronze Age through to the Hellenistic period, including the entire span of Greek occupation.

Approximately 90 burial mounds remain unexcavated at Apollonia (Davis et al., 2007) and thus the degree that our sample is representative of Apollonia’s population is unknown.

Corinth, Greece The Corinthian skeletal series represents the colonial mother-city and our “Greek” ancestry sample. The Corinthian skeletal material comes from several archaeological localities. Although we analyzed all known individuals that date to the pre-colonial and colonial periods at Corinth, our small sample (n585) may not accurately represent the ancient Corinthian population. Additionally, individuals dating to the colonial period were included in this article despite the fact that they may not be genetically Greek. Their inclusion is still appropriate because Corinthians are thought to have settled Apollonia in waves. Thus, any individuals living in Corinth during the colonial period could have contributed to the gene pool at Apollonia, regardless of whether or not they were purely of Corinthian or Greek ancestry. Interestingly, results of analyses including only those Corinthians who date to the prehistoric period (not reported on here) showed equivalent results to those presented below.

Lofkënd, Albania The skeletal series from Lofkënd represents the precolonial Illyrian population in this paper. Lofkënd lies roughly 15 miles east of Apollonia (Papadopoulos, 2006), and is the only site from central Albania with skeletal material that can be compared to Apollonia (Schepartz, 2010). The burial mound at Lofkënd was fully excavated and all individuals were considered in our analyses. Lofkënd largely predates colonization of the region (spanning the late Bronze through Early Iron Ages), but some grave goods at Lofkënd suggest that Lofkënd and Apollonia were contemporaneous for a short time (Papadopoulos, 2006; Papadopoulos et al., 2007).

Of the 227 individuals analyzed from Apollonia, 116 individuals dated to the Late Bronze Age through Hellenistic periods and had adult dentitions and crania with at least one nonmetric trait available for analysis. Of the 85 individuals from Corinth and 143 individuals from Lofkënd originally examined, 69 and 108 individuals, respectively, could be included here. Raw data for all skeletal samples used in this article and the age and sex distributions for each site have been published and are available for comparative use (see McIlvaine, 2012).


Methods We estimate patterns of biological affinity between the Illyrians and Greeks primarily using dental nonmetric data in conjunction with pseudo-Mahalanobis’ analysis, with both dental and cranial nonmetric data included in some analyses, as specified below (Mahalanobis, 1936; Konigsberg, 1990).

To test whether Illyrian and Greek individuals were buried together at Apollonia, we first employed dental nonmetric data. To confirm our results, given small sample sizes and missing data, we then examined a combination of dental and cranial nonmetric traits.

Due to the discrete nature of the data, we used “pseudo” Mahalanobis’ D² to measure biological distance, or dissimilarity, between sample populations. “Pseudo” D² accounts for correlations between phenotypic traits to avoid over-representing variation from characters that co-occur (Konigsberg, 1990; Scott andTurner, 1997; Edgar, 2004; Irish and Konigsberg, 2007).

All observable individuals from each population (Illyrian, Corinthian, and colonial Apollonian) were included in these tests.


Discussion The results here can be explained in three ways: (1) as the result of sample bias or methodological issues, (2) as a biologically meaningful indicator of close, shared ancestry between Greeks and Illyrians, or (3) as a combination of the previous two explanations. Each interpretation will be discussed in turn below.

Sample bias and methodological considerations The results of the biodistance studies described above occasionally contradict one another. These contradictions may represent a methodological issue with applying biological distance analyses to small samples. Three possible sources of error are identified: (1) small sample sizes, (2) the possible inability of the traits used here to capture variation among populations, and (3) heterogeneous skeletal samples.

Our results indicate that only small phenotypic differences exist between geographically adjacent Greece and Illyria. Thus, future research should attempt to expand the sample sizes used in our analyses. These sampling issues, along with a relatively homogenous phenotypic population throughout the Balkans, may be responsible for the fact that both Apollonians and many Greeks classify as Illyrian when the complete Illyrian sample isused in the creation of the logistic regression. Although the observed low biological distances among populations and the resultant high error rates in distinguishing between them suggest phenotypic similarity among these populations, it is also possible that the high errorrates indicate that the use of biological distance methods are inappropriate for small, fragmentary archaeological populations (Cheverud, 1988; Stojanowski and Schillaci, 2006).

Perhaps future researchers will identify new traits that can more accurately discriminate between the Greeks and Illyrians, or larger sample sizes will allow us to test the patterns we observed here with different trait combinations and larger numbers of variables. As with any bioarchaeological sample, the skeletal series in this analysis may not be fully representative of the Illyrian and Corinthian populations.


Shared ancestry and migration Despite possible complications, analysis of the current data does reveal patterns in biological variation among Apollonian, Corinthian, and southern Illyrian samples. All of the statistical analyses identified a general pattern of phenotypic similarity between post-colonial Apollonians and the prehistoric Illyrian samples. Although the following interpretations should be considered tentative due to issues with sample size and high error rates for some analyses, it is likely that the pattern of variation observed among colonial Apollonians, Corinthians, and the Illyrians, whereby biological similarity is demonstrated between the populations, is due to shared ancestry and long-term migration in the Balkan Peninsula.

Furthermore, colonial individuals buried in the Apollonian cemetery complex show more genetic similarity to the Illyrians from Lofkënd and prehistoric Apollonia than to the Corinthians. These data indicate that the population at Apollonia was primarily composed of Illyrians, with less genetic contribution from Corinthian colonists—as would be expected if only 200 Greeks initially colonized Apollonia (Hammond, 1992; Wilkes, 1992; Cabanes, 2008). The pattern remained constant in all tests,even when the Illyrian sample size was reduced to match the Corinthian sample size.

Although biological distances were smallest between the colonial Apollonians and Illyrians, biological distances were also small between the Corinthian and Illyrian populations in many analyses. Shared ancestry and long-term gene flow in the Balkan Peninsula may be responsible for the observed distribution of phenotypic characteristics. Our interpretation is consistent with the modern genetic record indicating that variation across the Balkans is clinally distributed with general homogeneity throughout the region (Mirabal et al., 2010). Y-chromosome evidence suggests that all Balkan populations, including the Greeks and Albanians, most likely migrated into the region in the Mesolithic period as foraging populations (Battaglia et al., 2009). Thus, genetic similarities among Greek and Illyrian populations likely stem from a shared ancestry, while phenotypic similarity may have been maintained through continuous migration in the form of traders in the region (Antonaccio,2001). In Illyria, the importation of Greek goods increased steadily into the Iron Age (Harding, 1992; Ceka, 2005), and at the time that the first Greek colonies in Illyria were founded—Epidamnus in 627 BC, Apollonia, ca. 600 BC—Greek pottery and other goods were frequently imported (Harding, 1992). Accompanying precolonial trade, interbreeding may have occurred among Greek and Illyrian populations. These data suggest that as a result of shared ancestry, close geographic proximity, and trade, accompanied by extensive gene flow among source populations, the southern Illyrians and Greeks shared certain biological features, specifically similar dental and cranial morphology, that came from a long history of biological interaction (Antonaccio, 2001; Mirabal et al., 2010).

Mortuary data also help to elucidate social dynamics of the colonies. Although Greek colonizers typically retained their tradition of building flat cemeteries, Greek colonies on Sicily, like Illyrian Apollonia, show departures in burial practice from their mother-cities (Shepherd, 1993). Few researchers have attempted to identify whether individuals buried adjacent to Greek colonies represent Greek colonists or locals. Those that have addressed this question show consistency with the findings of our article in that the majority of individuals are identified as locals (Stallo, 2007; Keenleyside et al.,2011). Our biodistance evidence for Illyrian population continuity at Apollonia contradicts Ceka’s (2005) suggestion that only Greeks were buried within the cemetery complex at Apollonia. Although the Greeks colonized the region, the majority of the general population may have remained primarily Illyrian in biological origin. The consistent pattern observed through logistic regression analysis suggests a strong Illyrian contribution to the gene pool at Apollonia. Finally, another possible interpretation of these data is that extensive interbreeding between Greeks and Illyrians makes it impossible to distinguish between groups (Galaty, 2002). In this case, few individuals from colonial Apollonia would represent a purely “Greek,” or “Illyrian” phenotypic signature. The colonial Apollonian skeletal series does represent the entire span of Greek occupation at Apollonia (roughly 500 years). Although most individuals from colonial Apollonia were classified by logistic regression analysis as Illyrian, regardless of how long after initial Greek colonization they lived, these results may be due to little phenotypic variability among individuals resulting from extensive gene flow between the Greeks and Illyrians. These data tentatively support the suggestion that the majority of individuals interred in the Apollonian cemetery complex were Illyrians, or the descendants of Greek and Illyrian intermarriages (Hammond, 1992; Wilkes,1992; Stallo, 2007).


NovoScriptorium: We get the impression that the researchers are more than ‘cautious’ to firmly announce that the Illyrian and Greek populations were almost identical in terms of phenotype, i.e., Anthropologically identical, something which clearly indicates common ancestry/ies. On top of that, taking in account that great affinity -almost identical everywhere- in the phenotypes of all ancient Balkan populations (something which is NOT valid for modern Balkan populations or even Medieval Balkan populations) is proved and accepted by modern Science, there are a few extra things to comment on.

We didn’t notice the same cautiousness when genomics scientists announced results about populations, migrations, etc, from the analysis of ONE or a few skeletons. Here we have analyses of hundreds of skeletons and still there is…cautiousness. We fully understand why we should always be cautious with similar results; but we should  always be cautious and not selectively, as this may imply other things (e.g. leading Science to ‘desired’ conclusions for political reasons).

It is written in the paper that only 200 Corinthians started the colonization of Apollonia. Similar conditions are mentioned for many other Greek colonies. Logic dictates that ‘they don’t just let you build a new city in a foreign land‘. This means 100% WAR, everywhere in the World, at any given human time. We must look at some other possibilities of how colonies started with just a few tens of people in ‘foreign lands’ – and without the use of armed force.

In our opinion, there are only two other alternatives besides war;

a) prior ethnic affinities – even distant in Time,

b) strong cultural and economic bonds prior to the colonization.

For those of us who don’t just ‘read’ the Greek Mythology, but ‘study’ it instead, it seems ‘brighter than the Sun’ that populations from the Greek peninsula expanded -not at a single time but in various waves over the millennia- over vast areas around them, including the Balkans, Western Europe, North Africa, Black Sea, Near East, Anatolia. We may not speak of ‘Greeks’ as in modern terms, even though the ancient Greeks referred to them as such – probably as an attempt for ethnic continuity to be denoted. There were many different names to describe the same population group over Time. One of them is the term ‘Greeks’. Another one, which we value as important for this commentary, is the term ‘Pelasgian’. Πελασγός derives from πέλας + γή or, according to another reference from Plato, from πελαργείν. Both terms indicate very interesting things. The first one refers to the ‘neigbouring population’ – in its plural form Πελασγοί. The second refers to ‘a population who migrates’, i.e. Πελαργοί.  Now, let’s add the initial myth about Πελασγός (Pelasgian).

Apollodoros delivers: «From Niovi and Zeus –she was the first mortal woman he mated with- were born Argos, as Acousilaos says, and Pelasgos, from whom the inhabitants of the Peloponnese were named Pelasgoi». «Acousilaos says that Argos was born of the earth (γηγενής) »

Argos is called ‘γηγενής’ (gegenes), meaning ‘born of the Earth’, indicating he was a native and not an immigrant from elsewhere (and this is what the ancient Greeks believed of their ancestries; that they were indigenous, locals from the very beginning – this obviously contradicts modern ideas of mass migrations from somewhere else,  replacement of local populations, etc.). The word ‘argos’ means ‘white’, ‘ablaze’, ‘resplendent’. The word ‘pelasgos’ means the ‘neighbouring’ or, if we accept another explanation of the word mentioned in Plato’s writings, means ‘migratory’. They were born from a common uterus (same mother) says the myth, and their cradle was the Peloponnese.

At some point, archaic  populations from the Peloponnese began to spread. These populations eventually took the name ‘Πελασγοί (Pelasgians)’. The cultural and ethnological affinity of the so called ‘prehistoric’ populations which archeologists happen to find in various sites, in Europe and the Mediterranean, is actually expected if one takes in consideration what the ‘prehistoric’ people themselves believed about their evolution.

The researchers have confirmed anthropological affinities between Illyrians and Greeks. It is not a surprise then that only a few Corinthians started a whole new reality for the Illyrian lands -and similar events took place in many other Greek colonies of the ‘Historical times’. People somehow knew that they were relatives. And they definitely shared some common cultural background.

One important thing that needs to be discussed is ‘why’, in the course of Time, cultural differences between Anthropologically similar populations apparently multiply.

Well, the ancient Greeks used the term ‘εκβαρβαρισμός’ and ‘βαρβαρίζοντες’ ‘barbarization’ and ‘barbarizing’ to denote populations relative to theirs but with weaker or stronger deviations from the culture of the ‘original cultural pond’. The term is already used in Homer to describe a linguistic barbarization of Anatolian populations (allies of the Trojans). Very few cultural differences are described to exist between Argives and allies vs Trojans and allies, while most of them -but not all- use the same language, too.

The very well known reality of the existence of different dialects in Greece itself in the ‘Historical times’ implies that this is a somehow ‘natural’ phenomenon. The further a population migrates to or the more the isolation of it becomes from the original Centre, the greater the possibility becomes for its ‘barbarization’, as the Greeks called it back then. Additionally, in cases where the migrant populations interbred with locals or migrants from elsewhere who moved in the same given area, the possibility of ‘barbarization’ becomes even greater – a certainty. For sure, the climate or any other local peculiarities strengthen the process.

Homer refers to Pelasgian inhabitants (probably denoting the oldest (?) migratory wave from the original Centre) at various places; Epirus and Thessaly, Crete, Thrace, Dardania, Asia Minor, among others.

Archaeological findings suggest that already from the Bronze Age (Mycenean culture), there were strong ties between the ‘Greeks’ and the Balkan populations (in our opinion, these relations must be very much older, at least from the Neolithic Age). In ancient Illyria, including the territory of modern Albania, this is equally true.

Hence, after all the above, for us, it comes as no surprise that Illyrians and Greeks co-founded new cities, interbred and produced a common culture, integrated in the ‘Greek World’; they already belonged to the same Anthropological and Cultural group. What, apparently, happened is that those much older bonds of theirs were ‘officially renewed’.


Research-Selection for NovoScriptorium: Philaretus Homerides

Comments for NovoScriptorium: Isidoros Aggelos

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