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Cranial Discrete Traits in a Byzantine Population and Eastern Mediterranean Population Movements

Article in Human Biology · November 2008 DOI: 10.3378/1534-6617-80.5.535 · Source: PubMed

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Francois X Ricaut Marc Waelkens Paul Sabatier University - Toulouse III KU Leuven

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The user has requested enhancement of the downloaded file. 1 Cranial Discrete Traits in a Byzantine Population and Eastern 2 Mediterranean Population Movements 3 4 5 f. x. ricaut1 and m. waelkens1,2 6 7 8 Abstract Since the beginning of the Holocene, the Anatolian region has been a crossroads for populations and civilizations from Europe, Asia, and 9 the Near to Middle East, with increasing interactions since the Bronze Age. 10 In this context, we examine cranial discrete traits from a Byzantine population 11 from southwest Turkey, excavated at the archeological site of Sagalassos; the 12 site displays human occupation since the 12th millennium b.p. To investigate [First Page] 13 the biological history of this population, we analyzed the frequency distribu- 14 tion of 17 cranial discrete traits from Sagalassos and 27 Eurasian and African [535], (1) 15 populations. Ward’s clustering procedure and multidimensional scaling anal- 16 yses of the standardized mean measure of divergence (MMDst), based on trait 17 frequencies, were used to represent the biological affinity between popula- Lines: 0 to 32 tions. Our results, considered within a large interpretive framework that takes 18 ——— 19 into account the idea that populations are dynamic entities affected by various influences through time and space, revealed different strata of the Sagalas- -1.40256pt PgVar 20 sos biological history. Indeed, beyond an expected biological affinity of the ——— 21 Sagalassos population with eastern Mediterranean populations, we also de- Normal Page 22 tected affinities with sub-Saharan and northern and central European popula- PgEnds: TEX 23 tions. We hypothesize that these affinity patterns in the Sagalassos biological 24 package are the traces of the major migratory events that affected southwest 25 Anatolia over the last millennia, as suggested from biological, archeological, [535], (1) 26 and historical data. 27 28 Since the beginning of the Neolithic period, the eastern part of the Mediterranean 29 basin and especially the Anatolian region, has been an area of interaction between 30 cultures and populations from Europe, northeast Africa, the Near to Middle East, 31 and the Eurasian steppe (Ammerman and Cavalli-Sforza 1984; Bellwood and Ren- 32 frew 2002; Richards et al. 2000). 33 The Neolithic and Chalcolithic periods have been relatively well documented 34 through numerous findings throughout southern Anatolia and upper Mesopotamia. 35 Nevertheless, the increasing number of archeological excavations and information 36

37 1Center for Archaeological Sciences, Katholieke Universiteit Leuven, Celestijnenlaan 200E, 3001 Heverlee, 38 Belgium. 39 2Sagalassos Archaeological Research Project, Katholieke Universiteit Leuven, Blijde Inkomststraat 21, 3000 Leuven, Belgium. 40 41 Human Biology, October 2008, v. 80, no. 5, pp. 535–564. 42 Copyright © 2008 Wayne State University Press, Detroit, Michigan 48201-1309 43 key words: cranial discrete traits, biodistance analysis, mean mea- 44 sure of divergence, byzantine population, sagalassos, anatolia.

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1 from ancient texts (e.g., Assyrian and Urartian annals, Hittite and Achaemenid 2 archives, and Greek, Roman, and Byzantine literary and epigraphic sources) make 3 the history of this area relatively better known from the second millennium b.c. 4 onward. The findings show that important cultural changes occurred in this region 5 since the Bronze Age because of increasing international contacts over a large area 6 (e.g., mainland Greece, the Aegean Islands, Anatolia, Syria, and Mesopotamia) 7 and development of often culturally overlapping and interacting civilizations (Ed- 8 wards et al. 2000; Sahoglu 2005). In this context, Anatolia occupied a key location 9 as a natural bridge between Europe and Asia and has been strongly influenced 10 through cultural, commercial, and population contacts from western, southern, 11 eastern, and northern neighboring regions since the Bronze Age: Hittite city-states 12 in central and south Anatolia (19th–12th century b.c.); the Arzawa federation in the 13 west (15th–12th century b.c.); Minoan (early 2nd millennium b.c. to 15th century 14 b.c.) and Mycenaean (15th–12th century b.c.) colonization of the west coast; the [536], (2) 15 Urartu kingdom in the east (10th– 8th century b.c.); Phrygian, Lydian, and Lycian 16 kingdoms in the west (12th–6th century b.c.); the Achaemenid Persian Empire, 17 including all of Anatolia (6th–4th century b.c.); the Roman republic and imperial Lines: 32 to 36 18 provinces (2nd century b.c.–5th century a.d.); Persian raids from the 3rd century ——— 19 a.d.; Byzantine civilization (6th–15th century a.d.); Arabic incursions from the 0.0pt PgVar 20 7th century a.d.; and the arrival of nomadic tribes from Central Asia from the 11th ——— 21 century a.d. (Boardman et al. 1984; Edwards et al. 2000; Mitchell 1994, 1995). Normal Page 22 The Anatolian population history is thus the result of complex processes involving PgEnds: TEX 23 different population groups for which the impact and contribution to the biological 24 diversity of the Anatolian population is relatively unknown. 25 Molecular genetic analysis of the modern Anatolian population has gen- [536], (2) 26 erally shown affinity with the Western Eurasian population group, despite the 27 presence of a substantial amount of lineages shared by Central Asian populations 28 and a few specific African lineages (Calafell et al. 1996; Cinniog¢lu et al. 2004; 29 Di Benedetto et al. 2001; Luis et al. 2004; Nasidze et al. 2004; Quintana-Murci 30 et al. 2004; Tambets et al. 2000). However, these results are limited by the fact 31 that the genetic structure of modern human populations has been influenced by 32 many historic, demographic, and genetic events (migration episodes, gene flow, 33 and genetic drift) that could have obscured the evolutionary history of Anatolian 34 populations (e.g., the presence of East Eurasian lineages in the current Turkish 35 population resulted from the immigration of nomadic groups from Central Asia in 36 the 11th century a.d.; Di Benedetto et al. 2001). In this context, analysis of ancient 37 populations appears to be a promising way to investigate population history and 38 to validate ethnogenetic hypotheses based on modern data. 39 Despite progress in genetic and biochemical analyses, the morphological 40 analysis of human remains is still the most frequently used method, and it is the 41 initial step to studying variability of ancient human populations. Several studies 42 based on morphological analysis from human skeletons from this geographic area 43 have been published, but they focused mainly on more ancient time periods (late 44 Paleolithic, Neolithic, Chalcolithic) and on issues that do not directly involve the

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1 origin and evolution of Anatolian populations (Bar-Yosef 1998; Bocquentin 2003; 2 Erdal 2008; Faerman et al. 2007; Hemphill 1998; Pearson 1999; Ullinger et al. 3 2005) and are rarely based on analysis of cranial discrete traits from Anatolians or 4 neighboring populations (Berry and Berry 1967, 1972; Hanihara and Ishida 2001a, 5 2001b; Parras 2004). 6 In this context the excavation of more than 50 tombs (11th–13th century 7 a.d.) at the (Hellenistic to Byzantine) Sagalassos archeological site (southwest- 8 ern Turkey) gives us a rare opportunity to address questions concerning the ori- 9 gin and evolution of this population up to the mid-Byzantine period. Within the 10 framework of the Sagalassos project, research is restricted to the period preceding 11 the capture of Constantinople by the Ottoman Turks in a.d. 1453. To fulfill this 12 purpose, we analyzed cranial discrete traits from this population and examined 13 biodistances with 27 Eurasian and African populations. Indeed, the analysis of 14 cranial discrete traits is one of the most efficient morphological approaches to [537], (3) 15 investigating interpopulation relationships (Berry and Berry 1967, 1972; Crubézy 16 1999; Donlon 2000; Hallgrimsson et al. 2004; Hanihara et al. 2003; Hanihara and 17 Ishida 2001a–d; Hauser and DeStefano 1989; Stefan and Chapman 2003; Sutter Lines: 36 to 53 18 and Mertz 2004), because discrete cranial traits provide biological distances sim- ——— 19 ilar to those found in genetic and other morphological analyses (Cavalli-Sforza et -0.42pt PgVar 20 al. 1994; Hanihara 2008; Hanihara et al. 2003; Howells 1995; Manica et al. 2007). ——— 21 However, even if there is a good correspondence between the results of biodis- Normal Page 22 tance studies using different kinds of data [see Relethford’s (2004) study based on PgEnds: TEX 23 blood cell polymorphisms, microsatellite DNA markers, and craniometric traits], 24 the population affinity patterns revealed from different biological markers, which 25 follow different evolutionary processes, may not always be concordant, as dis- [537], (3) 26 cussed by Keita (2004) and Keita et al. (2004) regarding the presence of the sub- 27 Saharan Benin sickle cell haplotype and Y-chromosome PN2 clade in southern 28 Europeans. 29 The purpose of this study is to document the cranial nonmetric traits of the 30 Byzantine Sagalassos population in order to investigate its biological history and 31 relationship with African and Eurasian populations in the context of the interac- 32 tions taking place in the Sagalassos region, but also on a larger scale in Anatolia 33 and the northeast Mediterranean, until Byzantine times. 34 35 The Archeological Site of Sagalassos: Historical Context 36 37 To evaluate the issues under investigation in this study, we need to provide 38 some fundamental information on how the Sagalassos region was affected and 39 therefore included in the network of regional population interactions during the 40 last millennia. 41 Human presence in the territory of Sagalassos (Figure 1) goes deep in time. 42 Indeed, the first traces of hunter-gatherers date back to the 12th millennium b.p., 43 and the first agricultural settlements, around the Burdur Lake, date to the 10th mil- 44 lennium b.p. (Waelkens et al. 1999). During the Bronze Age, territorial chiefdoms

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1 2 3 4 5 6 7 8 9 10 11 12 13 14 [538], (4) 15 16 17 Lines: 53 to 60 18 ——— 19 Figure 1. Location of the Sagalassos archeological site and comparative populations used in this study. 1, Pounbury (U.K.); 2, France; 3, Germany; 4, Scandinavia; 5, Italy; 6, Greeks; 7, -0.686pt PgVar 20 Eastern Europe; 8, Russia; 9, Turkey/Cyprus; 10, Naqada (Egypt); 11, Gizeh (Egypt); 12, ——— 21 Kerma (Egypt); 13, Somalia; 14, Tanzania; 15, Gabon; 16, Northwest India; 17, Tagars; Normal Page 22 18, Buryats; 19, Kazakhs; 20, Afghanistan; 21, Mongolians; 22, Northern Chinese; 23, PgEnds: TEX 23 Japanese; 24, Neolithic Baikalians; 25, Yakuts; 26, Evenks; 27, Asian Eskimos. 24 25 [538], (4) 26 developed in the region, whereas Sagalassos itself was most probably not yet oc- 27 cupied, despite evidence of agricultural activities near Sagalassos (Kaniewski et 28 al. 2008). This may have changed by the 14th century b.c., when the mountain site 29 of Salawassa (possibly to be identified with the later Sagalassos) was mentioned 30 in Hittite documents. 31 At the beginning of the 2nd millennium b.c. Indo-European tribes from 32 Transcaucasia occupied Anatolia: the Hittites in central Anatolia and the Luwiens 33 in southern and southwestern Anatolia. The territory of Sagalassos (and Sagalas- 34 sos itself) was included in the Luwien states but was located near the border with 35 the Hittite territories. 36 During the middle Bronze Age the Luwien populations were strongly in- 37 fluenced by the Minoan civilization (Crete), which colonized some part of the 38 Anatolian coast (e.g., north and east of the Dodecanese Islands). At the end of 39 the Minoan civilization (around the 15th century b.c.), the Mycenaeans (Greece) 40 took over the Minoan Empire (including the settlement areas). Mycenaean control 41 persisted until the 12th century b.c., when both the Mycenaean and Hittite Em- 42 pires collapsed. Then, the Phrygians, probably coming from Thrace, conquered 43 the Hittite Empire. On the Anatolian coast different kingdoms (e.g., Lydie, Lycie) 44 appeared with a Greek and Luwien admixed population. Greek settlers colonized

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1 the Anatolian coast from the northwest (Ilion-Troy, Lesbos, Assos, etc.) to the 2 central part of the occidental coast (Ephese, Milete, Priene, etc.). These settlers 3 probably reached the Pamphylienne coast (between Antalya and Alanya on the 4 southern coast of Anatolia), where several cities claim Greek origin (Aspendos, 5 Side, Perge) and from which Greek influence spread inland to Pisidia around the 6 5th century b.c. (Waelkens 2004). Pisidia, was part of the Phrygian and latter Ly- 7 dien Empires until it was conquered by the Persians in the 6th century b.c. 8 From the 6th to the 4th century b.c., Anatolia was included in the Persian 9 Achaemenid Empire. Persian aristocracy acquired large domains, and Persian pop- 10 ulations settled in the northern and central parts of the Anatolian plateau—just on 11 the northern border of the Sagalassos territory. In fact, Persian influence was still 12 strong during the Roman Empire, as Persian names appeared in Sagalassos. 13 Finally, Sagalassos appears in the limelight of history in 333 b.c., when the 14 town and the region of Pisidia were conquered by Alexander the Great. There- [539], (5) 15 after, Pisidia witnessed a sequence of Hellenistic kings. Greek culture diffused 16 throughout Anatolia through the influence of two Greek dynasties, the Ptolemaic 17 and Seleucid dynasties, which dominated Egypt and Syria-Mesopotamia-Anatolia Lines: 60 to 74 18 area, respectively. Furthermore, the Seleucids installed settlers from Macedonia in ——— 19 northern Pisidia, particularly in Sagalassos and in the Sagalassos territory (Wael- -0.50598pt PgVar 20 kens 2004). When the Attalids bequeathed their kingdom to Rome, the region of ——— 21 Pisidia was included in the republican province of Asia and later Cilicia. In 25 Normal Page 22 b.c., Rome decided to incorporate the region into its empire. The Pax Romana in- PgEnds: TEX 23 troduced by the soldiers of Augustus would last for centuries. During the Roman 24 domination, thousands of colonists from central Italy settled in the region of Pi- 25 sidia, which was close to the largest Roman colony (Antioch) in Anatolia (Mitchell [539], (5) 26 1994). During the following centuries, Roman colonists married Pisidian aristo- 27 crats, who themselves became Roman citizens (Mitchell and Waelkens 1998). 28 Sagalassos seems to have remained prosperous for centuries, but its glory 29 was shattered by an earthquake in the early sixth century a.d. Afterward, the set- 30 tlement may have been reduced to the status of a village. After the middle of the 31 seventh century a.d., a dramatic sequence of epidemics, Arab raids, and another 32 major earthquake took its toll (Waelkens 2002). Nevertheless, the last three years 33 of excavations allowed the discovery of three Byzantine villages, one of which 34 had been fortified and settled from the 8th to the 12th or 13th century a.d. Around 35 the 11th century a.d. Anatolia was invaded by nomadic tribes from Central Asia 36 (Seljuk Turk), and their presence is evident in the Sagalassos territory from at least 37 the 13th century (Vanhaverbeke et al. 2005). Finally, the Mongol invasion of the 38 13th century led to the division of the Seljuk kingdom, and at the end of the 13th 39 century, the Ottoman Empire overlapping three continents was created. 40 41 Materials and Methods 42 43 Samples. The archeological site of Sagalassos (southwestern Turkey) has been 44 the subject of an interdisciplinary research project coordinated by the Katholieke

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1 Universiteit Leuven since 1986 (Sagalassos Archaeological Research Project, 2 http://www.sagalassos.be). The Hellenistic to Byzantine site is located in the Tau- 3 rus Mountains of southwestern Turkey, 7 km from the modern village of Aglasun,˘ 4 in the ancient region of Pisidia (see Figure 1). The excavation of this site yielded 5 a city with monumental architecture and a necropolis (dated to a.d. 600–1200). 6 However, most of the buildings date from the 1st century b.c. to the 6th century 7 a.d. (Waelkens et al. 1999, 2006). 8 The burial site used in this study is located around a Christian church, and 9 recent archeological campaigns have provided the excavation of more than 50 10 graves containing skeletal remains of 57 individuals (Waelkens et al. 2006). The 11 deceased were all buried in pits, in some cases surrounded by lining stones, and 12 no clear indication for the use of coffins was observed. Funerary artifacts (bronze 13 crosses) and practices (east-west orientation of all the bodies) indicate that all the 14 individuals were buried according to the Christian tradition. Furthermore, the few [540], (6) 15 goods associated with the graves (in total, three earrings, some glass bracelets, 16 eight coins, and seven bronze crosses) and the simplicity of the tombs reflect a low 17 social status of the buried individuals. All the graves have been dated to the 11th– Lines: 74 to 117 18 13th century a.d. by AMS carbon-14 dating of human bones (11th–13th century ——— 19 cal a.d.) and/or from the stratigraphic sequencing coupled with the general histor- -0.12999pt PgVar 20 ical reconstruction of the site. The similarity in the funeral practices indicates that ——— 21 all these individuals shared a similar cultural package and probably belonged to Short Page 22 the same population group. Cranial discrete traits were collected only from adult PgEnds: TEX 23 individuals (37 in total) without cranial deformation. Age at death was primarily 24 estimated using evidence of fused sphenooccipital sychondrosis for the adults, as 25 well as epiphyseal fusion (White and Folkens 2005). The sex of the skeletons was [540], (6) 26 established according to standard anthropological techniques (White and Folkens 27 2005). 28 29 Discrete Traits. For this study, 35 nonmetric (i.e., epigenetic) cranial and man- 30 dibular traits were recorded for 37 adults buried at Sagalassos. The description of 31 each discrete trait examined in this study is described by Hauser and DeStefano 32 (1989). Although observations were made for each side in bilateral traits, for the 33 scoring procedure we calculated the trait frequencies using the individual count 34 method (Korey 1980; Turner and Scott 1977). A trait was scored as present if the 35 trait was present on either the left or the right side. The absence of side prefer- 36 ence in the case of unilateral expression has been demonstrated by several studies 37 (Cossedu et al. 1979; Hanihara and Ishida 2001a–d). Moreover, the scoring of 38 graded traits was converted to present or absent because the presence/absence form 39 of discrete trait scoring yielded the higher heritability estimate (Carson 2006). To 40 avoid interobserver error, the same person (F. X. Ricaut) collected all the cranial 41 and mandibular discrete traits used in this study. 42 The relatively poor preservation of the 37 skeletons did not allow us to 43 determine the sex in every case (only 54% of the individuals could be sexed). 44

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1 Table 1. Discrete Trait Frequencies for the Sagalassos Population for the 17 Nonmetric 2 Cranial Traits Used in the Biodistance Analysis 3 Cranial Trait Frequency Trait Description 4 5 Ossicle at lambda 3/21 (0.1428) Hauser and DeStefano (1989: 84–97) Ossicle at asterion 4/20 (0.20) Hauser and DeStefano (1989: 196–199) 6 Parietal notch bone 6/19 (0.315) Hauser and DeStefano (1989: 207–210) 7 Biasterionic suture 1/20 (0.05) Hauser and DeStefano (1989: 194–196) 8 Metopic suture 0/19 (0) Hauser and DeStefano (1989: 41–44) 9 Occipitomastoid bone 3/20 (0.15) Hauser and DeStefano (1989: 196) 10 Precondylar tubercule 4/21 (0.1904) Hauser and DeStefano (1989: 134–136) Condylius tertius 0/20 (0) Hauser and DeStefano (1989: 134–136) 11 Hypoglossal canal bridging 10/21 (0.476) Hauser and DeStefano (1989: 120–125) 12 Tympanic dehiscence 3/19 (0.1578) Hauser and DeStefano (1989: 143–147) 13 Patent condylar canal 11/19 (0.5789) Hauser and DeStefano (1989: 114–116) 14 Accessory infraorbital foramen 4/18 (0.222) Hauser and DeStefano (1989: 70–74) [541], (7) 15 Os japonicum 0/19 (0) Hauser and DeStefano (1989: 222–224) Jugular foramen bridging 2/19 (0.1052) Hauser and DeStefano (1989: 130) 16 Auditory exostosis 0/21 (0) Hauser and DeStefano (1989: 186–187) 17 Mylohyoid bridge 4/31 (0.1904) Hauser and DeStefano (1989: 234–236) Lines: 117 to 175 18 Accessory mental foramen 1/32 (0.03125) Hauser and DeStefano (1989: 230–233) ——— 19 -1.45944pt PgVar 20 ——— 21 Short Page 22 Moreover, the skulls were too fragmented to score all 35 cranial and mandibu- PgEnds: TEX 23 lar discrete traits. Consequently, investigation for significant sex association and 24 intertrait association was not systematically possible. Following the procedure of 25 several previous studies (Hanihara et al. 2003; Sutter and Mertz 2004), we pooled [541], (7) 26 the sexes in this study. Furthermore, as suggested by Hanihara et al. (2003), the 27 average intertrait correlation in human populations is relatively low (0.08) and the 28 patterns of geographic variation tend to be different from trait to trait, suggesting 29 the more or less independent expression of cranial discrete traits. 30 Of the 35 discrete traits examined from the Sagalassos specimens only those 31 shared with the comparison populations were selected for biodistance analysis. 32 The final trait list consists of 17 traits that are comparable to those analyzed in other 33 studies (Crubézy 1999; Hallgrimsson et al. 2004; Hanihara et al. 2003; Hanihara 34 and Ishida 2001a–d; Hauser and DeStefano 1989; Sutter and Mertz 2004). The 17 35 nonmetric cranial trait frequencies are presented in Table 1 and form the basis of 36 the distance analysis. 37 38 Populations Used for Comparative Analyses. To investigate the level of bi- 39 ological affinity of the Sagalassos population with the largest number of popu- 40 lations, we used the discrete trait data from 27 recent, historic, and prehistoric 41 African and Eurasian populations published by Hanihara and Ishida (2001a–d) 42 (Table 2). Few additional studies based on cranial discrete traits analysis have been 43 published for eastern Mediterranean populations (five in total): Greece, Cyprus, 44

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1 Table 2. Comparison Populations 2 Population a Period 3 4 Europe 5 1. Pounbury (UK) Late Roman 2. France Recent 6 3. Germany Recent 7 4. Scandinavia Recent 8 5. Italyb Recent 6. Greekb Ancient and recent 9 b 10 7. Eastern Europe Recent 8. Russia Recent 11 Middle East 12 9. Turkey/Cyprusb Recent 13 Sagalassos (Anatolia)b,c 7–13th century a.d. Northeast Africa [542], (8) 14 b 15 10. Naqada (Egypt) 3–2nd millennium b.c. 11. Gizeh (Egypt)b 7–4th century b.c. 16 12. Kerma (Egypt)b 20–16th century b.c. 17 Sub-Sahara Lines: 175 to 175 18 13. Somalia Recent ——— 19 14. Tanzania Recent 1.18513pt PgVar 20 15. Gabon Recent India ——— 21 16. Northwest India Recent Normal Page 22 Central Asia PgEnds: TEX 23 17. Tagarsb Iron Age 18. Buryats Recent 24 b 25 19. Kazakhs Recent [542], (8) 20. Afghanistan Recent 26 East Asia 27 21. Mongolians Recent 28 22. Northern Chinese Recent 29 23. Japanese Recent 30 Northeast Asia 24. Neolithic Baikaliansb Recent 31 25. Yakuts Recent 32 26. Evenks Iron Age 33 27. Asian Eskimos Recent 34 a. For a more detailed description of these populations, see Hanihara et al. (2003) and Hanihara and 35 Ishida (2001a–d). 36 b. Populations included in the regional data set. 37 c. This study. 38 39 40 Syria, ancient Palestine, and Egypt (Berry and Berry 1967; Parras 2004). These 41 data could not be used in our study because it would have reduced the power of 42 our statistical analysis by decreasing the number of populations and discrete traits 43 used; these five additional populations have been analyzed for a different number 44 of discrete traits (6 and 30 discrete traits used by Parras (2004) and Berry and

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1 Berry (1967), respectively) and had only 2 and 10 discrete traits in common with 2 those analyzed from the Sagalassos population and those used by Hanihara and 3 Ishida (2001a, 2001b). Consequently, to ensure a robust statistical analysis, we 4 considered only the populations from Hanihara and Ishida (2001a, 2001b) because 5 they allowed us to work from the largest number of cranial discrete traits (17) and 6 populations (27). 7 The comparison population data set used in this study was composed of 8 27 populations, which were pooled into two data sets to perform analyses on the 9 global and regional scales (see Table 2). The geographic location of these popu- 10 lations is shown in Figure 1, and more detailed information regarding their arche- 11 ological, geographic, and ethnological contexts, as well as their discrete trait fre- 12 quency can be found in Hanihara and Ishida (2001a–d) and Hanihara et al. (2003). 13 14 Statistical Methods. To estimate the biological distance between populations, [543], (9) 15 we used a method that calculates the standardized mean measure of divergence 16 (MMDst), which is more appropriate than the unstandardized MMD when the 17 number of observations for some traits is small or the sample sizes different Lines: 175 to 208 18 (Sjøvold 1973). This method (mean measure of divergence) for estimating the ——— 19 biological distance between each pair of samples is currently the method most 0.84804pt PgVar 20 used in the literature, and thus it allows a comparison of the results of different ——— 21 studies. Normal Page 22 The first step in our analysis was to stabilize variances by using an arcsine PgEnds: TEX 23 transformation of the trait frequencies (Green and Suchey 1976). This transforma- 24 tion allows us to use small samples as well as traits with a frequency less than 5% 25 or greater than 95%. The arcsine transformation is calculated using the equation [543], (9) 26     1 − 2k 1 − 2(k + 1) 27 θ = 0.5 arcsin + 0.5 arcsin , (1) 28 n + 1 n + 1 29 30 where k is the score of the considered trait and n is the number of subjects exam- 31 ined for this trait from the population. When k equaled 0, we applied the Bartlett = 32 correction such that k/n 1/4n. 33 We used the arcsine-transformed trait frequencies to calculate the mean mea- 34 sure of divergence (MMD):   35 N 36 2 1 1 (θ i − θ i ) − + 1 2 1 1 37 n i + n i + = i=1 1 2 2 2 , 38 MMD N (2) 39 40 where N is the number of traits considered, n1i and n2i are the number of individuals 41 examined for the trait in populations 1 and 2, respectively, and θ1i and θ2i are the 42 transformed frequencies of the ith trait in populations 1 and 2, respectively. The 43 MMD can have negative values (Sjøvold 1973), and this occurs when there is very 44 little difference or no difference in the frequencies of the traits considered.

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1 The variance of the MMD is 2 2 3 Var =   . (3) MMD N 2 4  1 1 N 2 + 5 1 1 n i + n i + 6 i=1 1 2 2 2 7 The standard deviation of the MMD is determined by 8 9 = ( )1/2. 10 SDMMD Var MMD (4) 11 Finally, the standardized MMD is calculated as 12 13 MMD = MMD/SD . (4) 14 st MMD [544], (10) 15 The MMDst distances are conventionally considered statistically significant if their 16 value is greater than 2 (de Souza and Houghton 1977). However, the interpretation 17 of statistical significances may lead to ambiguous statements; by assuming that no Lines: 208 to 288 18 statistical significant difference is found between populations (value less than 2), ——— 19 it is implied that there is no difference between them (Armstrong 2007). To avoid 11.43619pt PgVar 20 ——— this misconception, we do not interpret the MMDst value in terms of absence or 21 presence of statistical significant difference between populations but rather as an Short Page 22 indicator of the degree of biological affinity between them. In addition, although PgEnds: TEX 23 we indicated that values greater than 2 are considered statistically significant, none 24 of the hypotheses and conclusions of our study will be drawn from the significance 25 [544], (10) of the MMD values alone; we will always consider archeological, historical, and 26 st biological evidence. 27 The matrix of MMD values is analyzed using two different methods— 28 st 29 Ward’s (1963) hierarchical clustering procedure and a nonmetric multidimen- 30 sional scaling (MDS) procedure (Kruskal 1964)—to provide two perspectives on 31 the patterning of intersample affinities. Ward’s clustering procedure minimizes 32 intracluster variation while maximizing intercluster variation. This analysis was 33 performed using the R statistical package (R Project 2005). The MDS analysis 34 of MMDst matrices was performed using the Xlstat software (http://www.xlstat 35 .com/en/home/). This procedure produces graphical representations that are eas- 36 ily interpreted and that more accurately reflect complex or non-Euclidean distance 37 matrices, as in the case of MMDst values (Kruskal and Wish 1984). 38 Furthermore, we performed the cluster analysis and MDS after pooling our 39 comparison population data sets in two different ways to make a comparison: 40 (1) on a global scale, by using the global data set, which includes 28 populations 41 (Table 2); and (2) on a regional scale, by using a regional data set that includes only 42 the most relevant populations to the region based on archeological and historical 43 data (11 populations, as described in Table 2). 44

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1 Table 3. Mean Measure of Divergence (MMD) and Its Standard Deviation (SD) and 2 Standardized MMD (MMDst) Values Based on 17 Nonmetric Traits Between Sagalassos 3 and the 27 Comparison Populations a 4 Population MMD SD MMDst 5 Mongolians 0.11 0.02 5.83 6 Baikalians 0.05 0.02 2.42 7 Yakuts 0.08 0.02 3.53 8 Northern Chinese 0.10 0.02 5.36 9 Afghanistan 0.05 0.02 2.15 10 Asian Eskimos 0.10 0.02 4.95 Evenks 0.08 0.02 4.32 11 Japanese 0.10 0.02 5.65 12 Buryats 0.13 0.02 6.75 13 Northwest India 0.05 0.02 2.59 14 Tagars 0.05 0.02 2.55 [545], (11) 15 Kazakhs 0.05 0.02 2.44 Turkey/Cyprus 0.02 0.02 0.98 16 Greeks 0.01 0.02 0.56 17 Eastern Europe 0.02 0.02 1.15 Lines: 288 to 300 18 France 0.03 0.02 1.24 ——— 19 Russia 0.01 0.1 4.95 -0.81514pt PgVar 20 Italy 0.01 0.02 1.09 Germany 0.02 0.02 1.02 ——— 21 Scandinavia 0.01 0.02 0.72 Short Page 22 Pounbury (U.K.) 0.01 0.03 1.48 PgEnds: TEX 23 Naqada (Egypt) 0.01 0.03 1.38 24 Gizeh (Egypt) 0.01 0.03 1.49 25 Kerma (Egypt) 0.01 0.03 1.46 [545], (11) Somalia 0.01 0.04 1.68 26 Tanzania 0.01 0.07 3.65 27 Gabon 0.01 0.04 1.93 28 a. Values greater than 2.00 are statistically significant at the 0.05 level. 29 30 31 32 Results 33 34 MMDst values calculated from 17 nonmetric traits between Sagalassos and 35 the 27 Eurasian populations are provided in Table 3. The matrix of MMDst values 36 between each pair of populations is not shown because of its unwieldy size (data 37 available on request from the authors). 38 An examination of the biodistances (Table 3) shows that the Sagalassos 39 population is more similar to West Eurasian and ancient northeast African pop- 40 ulations than to Central and East Eurasian populations. The closest populations 41 to Sagalassos are from Greece, Cyprus and Turkey, Germany, and Scandinavia, 42 followed by the other European and ancient northeast African (ancient Egyptian 43 and Sudanese) populations and then by the Central and East Eurasians and the 44

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1 2 3 4 5 6 7 8 9 10 11 12 13 14 [546], (12) 15 16 17 Lines: 300 to 314 18 ——— 19 -6.686pt PgVar 20 Figure 2. Multidimensional scaling plot based on the standardized mean measure of divergence val- ——— 21 ues showing relationships between the 28 populations from the global data set used in Normal Page 22 this study. The stress value for the MDS plot is 0.180. European populations (black filled PgEnds: TEX 23 squares): POU, Pounbury (U.K.); FRA, France; GER, Germany; SCA, Scandinavia; ITA, 24 Italy; GR, Greece; EAE, Eastern Europe; RUS, Russia. Near Eastern population (gray filled 25 square): TRK, Turkey/Cyprus. Indian population (white filled square): NWI, northwest In- [546], (12) dia. Northeast Asians (black filled circles): BAI, Baikalians; YAK, Yakuts; EKV, Evenks; 26 ESK, Asian Eskimos. East Asians (gray filled circles): MON, Mongolians; NCH, northern 27 Chinese; JPN, Japanese. Central Asians (white filled circles): TAG, Tagars; BUR, Bury- 28 ats; KAZ, Kazakhs; AFG, Afghanistan. Africans (gray filled diamonds): NAQ, Naqada 29 (Egypt); GIZ, Gizeh (Egypt); KER, Kerma (Egypt); SOM, Somalia; TNZ, Tanzania; GAB, 30 Gabon. Sagalassos population (black filled triangle): SAG, Sagalassos. 31 32 33 sub-Saharan Tanzanian population. Intriguingly, the closeness of the Sagalassos 34 population to Germans and Scandinavians was unexpected, but more surprising 35 and less obviously explainable were the MMDst values from Gabon and Somalia, 36 which show some similarity with the Sagalassos population, yet the MMDst scores 37 are nearly significant (Gabon, 1.93; and Somalia, 1.68; see Table 3). 38 The MDS representation of the global data set of 28 populations (Figure 39 2) shows roughly three main population clusters: (1) Central, Northeast, and East 40 Eurasian populations, which are found in the top left; (2) West Eurasian and an- 41 cient Egyptian and Sudanese populations in the lower part; and (3) recent sub- 42 Saharan populations in the top right. The Sagalassos population clusters with the 43 second group and is most closely related to Greek, Cypriot/Turkish, and Scandi- 44 navian populations.

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1 2 3 4 5 6 7 8 9 10 11 12 13 14 [547], (13) 15 16 17 Lines: 314 to 333 18 ——— 19 -6.686pt PgVar 20 ——— 21 Normal Page 22 PgEnds: TEX 23 24 25 Figure 3. Dendrogram of Ward’s hierarchical cluster analysis based on the MMDst distance ma- [547], (13) 26 trix obtained from the global data set of 28 populations. GER, Germany; EAE, Eastern Europe; FRA, France; ITA, Italy; POU, Pounbury (U.K.); GIZ, Gizeh (Egypt); NAQ, 27 Naqada (Egypt); KER, Kerma (Egypt); TRK, Turkey/Cyprus; GR, Greece; SCA, Scandi- 28 navia; GAB, Gabon; SOM, Somalia; TNZ, Tanzania; EKV, Evenks; ESK, Asian Eskimos; 29 NCH, northern Chinese; JPN, Japanese; MON, Mongolians; YAK, Yakuts; BUR, Buryats; 30 NWI, northwest India; RUS, Russia; TAG, Tagars; KAZ, Kazakhs; BAI, Baikalians; AFG, 31 Afghanistan. 32 33 34 The dendrogram produced by Ward’s clustering procedure for the global 35 data set is shown in Figure 3 and provides a relatively similar representation of 36 the MMDst distance matrix than that provide by the MDS analysis. The popula- 37 tions clearly fall into two groups. The first main group can be broken down into 38 two subgroups: (1) all the recent sub-Saharan populations and (2) mainly Cen- 39 tral, East, and Northeast Eurasians. West Eurasians form the second main group, 40 which is also subdivided into two subgroups. One of these subgroups includes 41 all the eastern Mediterranean populations (three ancient Egyptian/Sudanese pop- 42 ulations from Naqada, Gizeh, and Kerma as well as the Cypriot/Turkish, Greek, 43 and Sagalassian populations) and the Scandinavian sample; the second subgroup 44 includes the other West Eurasian populations.

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1 2 3 4 5 6 7 8 9 10 11 12 13 14 [548], (14) 15 16 17 Lines: 333 to 333 18 ——— 19 0.314pt PgVar 20 ——— 21 Normal Page 22 PgEnds: TEX 23 24 Figure 4. Multidimensional scaling plot based on standardized mean measure of divergence values showing relationships between the 11 regional populations used in this study. The stress 25 value for the MDS plot is 0.095. European populations (black filled squares): ITA, Italy; [548], (14) 26 GR, Greece; EAE, Eastern Europe. Near Eastern population (gray filled square): TRK, 27 Turkey/Cyprus. Northeast Asians (black filled circle): BAI, Baikalians. Central Asians 28 (white filled circles): TAG, Tagars; KAZ, Kazakhs. Africans (gray filled diamonds): NAQ, 29 Naqada (Egypt); GIZ, Gizeh (Egypt); KER, Kerma (Egypt). Sagalassos population (black filled triangle): SAG, Sagalassos. 30 31 32 33 Cluster analysis and MDS representations from the regional data set (Fig- 34 ure 4 and 5) are in agreement with the results obtained from the global data set. 35 However, the regional populations show no unexpected affinity pattern; rather, as 36 the geographic distance increases between regional populations, the dissimilarity 37 in cranial discrete traits also increases. Consequently, as seen in Figures 4 and 38 5, the regional populations form two discernible groups. The Central Eurasian 39 populations cluster together in the first group. The second group includes the 40 West Eurasians and northeastern African populations and is divided into three 41 subgroups. The first subgroup includes the geographically closest populations to 42 Sagalassos: Cypriots/Turkish, Greeks, and the ancient Egyptian population from 43 Gizeh (located in the northern Nile delta). The second and third subgroups include 44 the last two northeastern African populations (located far south of the Mediter-

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1 2 3 4 5 6 7 8 9 10 11 12 13 14 [549], (15) 15 16 17 Lines: 333 to 340 18 ——— 19 10.314pt PgVar 20 ——— 21 Normal Page 22 PgEnds: TEX 23 24 25 [549], (15) 26 Figure 5. Dendrogram of Ward’s hierarchical cluster analysis based on the MMDst distance matrix obtained from the regional data set of 11 populations. KAZ, Kazakhs; BAI, Baikalians; 27 TAG, Tagars; GIZ, Gizeh (Egypt); GR, Greece; TRK, Turkey/Cyprus; EAE, Eastern Eu- 28 rope; ITA, Italy; NAQ, Naqada (Egypt); KER, Kerma (Egypt). 29 30 31 32 33 ranean coast) and two European populations (Italy and Eastern Europeans), re- 34 spectively. 35 Finally, a detailed review of the different statistical tests (MMDst; MDS and 36 Ward clustering) shows that the unexpected biological proximity of some northern 37 and central European and sub-Saharan populations to the Sagalassos population is 38 not supported to the same significance. Indeed, as seen by the MMDst values dis- 39 played in Table 3, Scandinavians and Germans (MMDst of 0.72 and 1.02, respec- 40 tively) present stronger affinity to Sagalassos than populations from Somalia and 41 Gabon, which have nearly significant MMDst values (1.68 and 1.93, respectively). 42 In addition, only the biological affinity between the Sagalassos and Scandinavian 43 populations suggested by the MMDst values (see Table 3) is preserved when all the 44 comparative populations are considered (see Figures 2 and 3). Nevertheless, we

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1 discuss the validity of such affinity patterns in the light of archeological, historical, 2 and biological data. 3 4 Discussion 5 6 Limitations and Biases. The results obtained in this study, as with many stud- 7 ies, are presented with a number of unavoidable caveats. In particular, the use of 8 morphological variants, which have an etiology based on a combination of envi- 9 ronmental and hereditary factors, to measure biological distances is problematic. 10 Furthermore, the analysis of a single biological trait (such as nonmetric traits), 11 which gives only a crude indication of evolutionary history (because gene and 12 biological trait frequencies are subject to random sources of variation; Cavalli- 13 Sforza et al. 1994) is an important limitation. Nevertheless, several studies have 14 shown that nonmetric traits (and craniometric data) offer an efficacious means to [550], (16) 15 investigate origin and evolution of human populations (Hanihara 2008; Hanihara 16 et al. 2003; Howells 1995). In addition, the pattern of interpopulation relationships 17 obtained in our study is similar to that obtained by Hanihara et al. (2003, Figures 1– Lines: 340 to 350 18 3), suggesting that analyses performed from the 17 cranial discrete traits selected ——— 19 in our study provide reliable results. -0.28598pt PgVar 20 This study also has limitations because cemetery samples do not typically ——— 21 conform to the standard definition of a population and biases are inevitable. One Normal Page 22 bias of our study is related to the relatively small number of skulls from Sagalassos PgEnds: TEX 23 analyzed for each discrete cranial trait (between 18 and 32 skulls per trait). More- 24 over, no more than 50 tombs have been excavated, and the skull sample probably 25 represents a subset of the total individuals buried between the 11th and 13th cen- [550], (16) 26 turies a.d. Consequently, there is a risk that the Sagalassos skull sample analyzed 27 in our study does not properly reflect the Sagalassos population living during this 28 period, and unfortunately we cannot evaluate the degree of discrepancy between 29 the Sagalassos archeological sample and the Sagalassos living population. Never- 30 theless, the Sagalassos population sample used in this study comes from a geo- 31 graphically (same burial place), temporally (11th–13th century a.d.), and cultur- 32 ally (same funeral practices) restricted area, which allows us to assume that our 33 sample represents a sequence of related individuals and thus can be equated to a 34 biological population (Konigsberg 1990). This has implications in terms of intr- 35 asite variation and biological history. First, as explained by Stefan and Chapman 36 (2003), if the sample encompasses several generations and can then be considered 37 a “sequence of related individuals,” the comparison between sites should not be 38 affected by intrasite variation, and therefore ancient populations can be consid- 39 ered statistically equivalent to a biological population. Second, when considering 40 the historical and demographic events that affected the Sagalassos area during the 41 last millennium, diachronic variation of the Sagalassian morphology might be ex- 42 pected over this period. Nevertheless, our population sample, with its relatively 43 small number of individuals from a restricted time period, did not allow us to 44 analyze Sagalassos diachronic variations.

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1 A final limitation of the study is due to the nature of the populations used for 2 comparative analysis. Indeed, the comparison populations do not cover the full ge- 3 ographic range involved in the interactions with Anatolia during the last millennia 4 [e.g., no data were available from the Middle East (especially Iran/Persia) or Tran- 5 scaucasia], and they come from different time periods. Moreover, samples from the 6 northeastern Mediterranean region, which is the area historically most implicated 7 in the interaction network with Anatolian populations, are lacking. Specifically, 8 we do not have cranial discrete trait data from all the different populations and re- 9 gions potentially involved (e.g., Macedonians, Mycenaeans from Greece, Minoans 10 from Crete, Phrygians from Thrace), and we do no know the degree of biological 11 differentiation between these geographically close populations. The first problem 12 can be solved only through additional archeological excavation and further bio- 13 logical analysis of the human remains. Nevertheless, for the second problem dif- 14 ferent studies have shown that biological affinity between populations, based on [551], (17) 15 cranial discrete traits, is shaped by geographic factors rather than by the tempo- 16 ral origins of these populations (e.g., Donlon 2000; Hanahira et al. 2003). These 17 studies revealed higher biological affinity between two populations from different Lines: 350 to 356 18 time periods in the same geographic area rather than a higher affinity between two ——— 19 contemporary populations from different geographic locations [with the exception -0.34999pt PgVar 20 of a few isolated populations (e.g., Ainu, Andaman), which can show a different ——— 21 pattern]. Thus the gradient of differentiation between populations appears to be Normal Page 22 more geographically than temporally based. These results indicate that, even if PgEnds: TEX 23 we do not know the biological features of the populations when they potentially 24 interacted with southwestern Anatolian populations, we can infer that, in the ab- 25 sence of major population replacement, ancient and modern populations from the [551], (17) 26 same location are biologically closer to each other than to any other populations. 27 Keeping these considerations in mind, and despite the limitations described, 28 our results add substantial information on the issues investigated in this study. 29 30 Sagalassos Affinity with Eastern Mediterranean Populations. The results 31 obtained in this study (see Table 3 and Figures 2–5) reveal that the Byzantine popu- 32 lation from Sagalassos clusters with the West Eurasian and northeast African pop- 33 ulations. In each of the analyses performed the Byzantine population is included 34 in the regional (e.g., eastern Mediterranean) biological variability, with the closest 35 affinity to populations from Greece and Cyprus/Turkey rather than to Italy and 36 ancient Egypt. This is consistent with most anthropological studies, which show a 37 positive correlation between geographic separation and biological (phenotypic or 38 genetic) distance among populations (Crawford 1998; Konigsberg 1990; Manica 39 et al. 2007; Relethford 2004). Nevertheless, a few populations in our study (Scan- 40 dinavians, Germans, Gabonese, and Somalians) did not fit this pattern; they are 41 geographically distant from Sagalassos but biologically close (these differences 42 are discussed in a later section). 43 Our results have several implications related to the historical, archeological, 44 and biological framework of interactions occurring in this geographic area during

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1 the last millenniums. First, our results (see Table 3 and Figures 2–5) show the 2 low biological affinity between the Sagalassos population and populations from 3 Central Asia (MMDst from 2.44 to 6.75). If our Central Asian population samples 4 are not too different from the 11th-century Oghuz Turks, then these results are in 5 agreement with historical records and genetic studies. Indeed, these studies show 6 that nomadic tribes from Central Asia (Oghuz Turks) have been documented in 7 the area between Mongolia and the Caspian Sea since the 9th century a.d. Under 8 the rule of the Seljuk dynasty, these tribes invaded Anatolia in 1071 (Roux 1984) 9 and probably facilitated a continuous migratory contact between Anatolia and its 10 Asian neighbors. This continuous immigration from Central Asia occurred at a 11 relatively low rate until the language barrier disappeared a few centuries later and 12 facilitated gene flow between linguistically related areas, even if the impact of 13 this process is still being debated (Berkman et al. 2008; Cinniog¢lu et al. 2004; Di 14 Benedetto et al. 2001). As expected, the Sagalassos population predates the period [552], (18) 15 of highest gene flow from Central Asia and does not show any biological affinity 16 through cranial discrete traits to Central Asian populations. 17 Second, all our results indicate a closer affinity of the Sagalassos population Lines: 356 to 365 18 to its closest geographic neighbors (Greek and Cypriot/Turkish populations) rather ——— 19 than to any other eastern Mediterranean populations used in this study (Italians and 0.0pt PgVar 20 ancient Egyptians). ——— 21 In the general context defined previously, the close relationship among the Normal Page 22 northeastern Mediterranean populations (e.g., Sagalassos, Greek, and Cypriot/ PgEnds: TEX 23 Turkish populations) is in agreement with (1) their geographic proximity (model 24 of isolation by distance), (2) a likely common shared ancestor dating back to at 25 least the early Holocene (see the later discussion regarding Sagalassos affinity with [552], (18) 26 sub-Saharan and northern and central European populations), and (3) increasing 27 interactions between these neighboring regions and populations since the Bronze 28 Age, as suggested by linguistic, archeological, and mythological studies (Edwards 29 et al. 2000; Sahoglu 2005; Vanhaverbeke and Waelkens 2003), which were accom- 30 panied by population movements and some degree of gene flow, as suggested by 31 recent genetic studies (Cinniog¢lu et al. 2004; Cruciani et al. 2007; Di Benedetto 32 et al. 2001; King et al. 2008; Tambets et al. 2000). However, it is not possible 33 from the population sample and the biological data used in this study to detect 34 which historical populations (e.g., Minoans, Mycenaeans, Macedonians) might 35 have contributed, and in what proportion, to the Sagalassos biological variabil- 36 ity. To summarize, the affinity among northeastern Mediterranean populations is 37 probably due to common shared ancestry and has been accentuated by subsequent 38 gene flow between these regions and populations. 39 Concerning the affinity of the Sagalassos population with ancient Egyptian 40 populations, different lines of evidence suggest that this affinity is probably very 41 old (dating back to at least the early Holocene) and is more likely due to com- 42 mon shared ancestry than to recent gene flow, or to convergent adaptation (see 43 the later discussion regarding the affinity between Sagalassos and sub-Saharan 44 populations).

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1 Another interesting result is the lower biological affinity of the Sagalassos 2 population with the Italian sample (MMDst = 1.09) compared to the Greek or 3 Cypriot/Turkish samples (see Table 3 and Figures 2 and 3), in agreement with 4 genetic studies (Cinniog¢lu et al. 2004; Cruciani et al. 2007; Richards et al. 2000; 5 Semino et al. 2004;). This raises the question of the impact of the thousands of 6 colonists from central Italy who settled in Pisidia, particularly in the Sagalassos 7 territory, during the Roman period (Mitchell and Waelkens 1998). From our results 8 we did not detect any strong affinity between the Sagalassian and Italian samples, 9 suggesting either that no major admixture events occurred between Roman set- 10 tlers and the ancestors of the Sagalassos Byzantine sample or that some degree of 11 admixture occurred but is unfortunately not detectable in our samples (sampling 12 bias) or by our methods. Considering the historical and archeological context, the 13 Roman colonists belonged to an upper class and married mainly Pisidia aristo- 14 crats. In contrast, our population sample from Sagalassos represents a low social [553], (19) 15 class, which is less likely to include any descendant of the admixed elite popu- 16 lation living during the Roman time. For the affinity between Sagalassos and the 17 Egyptian populations, we can postulate, then, that the affinity pattern between the Lines: 365 to 369 18 Sagalassos and Italian samples may have resulted from common shared ancestry ——— 19 rather than from recent gene flow from Italian populations. -0.34999pt PgVar 20 ——— 21 Sagalassos Affinity with Sub-Saharan and Northern and Central European Normal Page 22 Populations. Finally, as noted previously, intriguing affinity patterns of the PgEnds: TEX 23 Sagalassos population have been detected without obvious explanations: on the 24 one hand, with two populations from northern and central Europe (Scandinavia 25 and Germany); and on the other hand, with two sub-Saharan populations (Soma- [553], (19) 26 lia and Gabon). Such an unexpected biological affinity between populations based 27 on cranial discrete traits has already been reported in the literature but without 28 clear hypotheses being suggested (Berry and Berry 1967; Donlon 2000; Hanihara 29 et al. 2003). Nevertheless, if we exclude potential statistical and methodological 30 errors, these unusual affinity patterns may have been caused by three different fac- 31 tors: convergent adaptation to local environment, gene flow, and common ancestry 32 (Gonzalez-Jose et al. 2003). 33 The first factor, convergent adaptation to local environment, is unlikely to be 34 the main cause of cranial discrete trait affinity of the Sagalassos population with 35 some northern and central Europeans and sub-Saharans, for three reasons. First, 36 it would be unlikely that local environmental pressures or any microevolutionary 37 mechanisms (e.g., mutation, drift, and selection) similarly occurred in populations 38 from Anatolia (Sagalassos), West Africa (Gabon), the Horn of Africa (Somalia), 39 and northern and central Europe (Germany and Scandinavia) to result in the affin- 40 ity patterns observed in this study. Second, with the exception of a few specific 41 morphological processes (i.e., gracilization; Lahr 1996), there is no evidence in 42 modern humans that environmental adaptation (through developmental plasticity 43 and climatic adaptation through natural selection) is one of the main factors that 44 shapes morphology and strongly affects cranial phenotypic traits (Gonzalez-Jose

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1 et al. 2003; Hanihara et al. 2003; Relethford 2002), which are known to respond 2 adaptively only to extreme environmental conditions (Roseman 2004; Roseman 3 and Weaver 2004). Third, even if we do not discard environmental adaptation as 4 a plausible factor, environmental conditions have always been variable, since at 5 least the late Pleistocene, in the Taurus Mountains of Anatolia, West Africa, the 6 Horn of Africa, and northern and central Europe (Bradley 1999), and it is unlikely 7 that different environmental conditions would have led to cranial trait affinity of 8 the population from the Taurus Mountains with those from the other regions. 9 The discussion related to the role played by the second (gene flow) and third 10 (common ancestry) factors in the observed population affinity pattern must take 11 into account the fact that these two factors are sometimes difficult to differentiate 12 (e.g., two populations have shared a common ancestor and experienced significant 13 gene flow between them after they diverged). In addition, “populations should be 14 viewed processually as dynamic entities over time and not static entities” (Keita [554], (20) 15 2004: 688); that is, population biological features may vary through time because 16 a population may have been affected by various influences. Last, population bi- 17 ological history suggested from different biological markers (e.g., morphologic, Lines: 369 to 377 18 classical polymorphisms, genetic markers) is not necessarily concordant, implying ——— 19 different levels of biological history (Keita 2004; Keita et al. 2004). 0.0pt PgVar 20 Keeping in mind these three elements, if we consider the affinity of the ——— 21 Sagalassos population with the sub-Saharan populations from Gabon and Soma- Normal Page 22 lia, a recent direct contact between these populations and regions probably can PgEnds: TEX 23 be excluded because they are separated by significant geographic distances. How- 24 ever, indirect contacts through geographically intermediary populations carrying 25 “sub-Saharan” biological features in the late Pleistocene–Holocene period are dis- [554], (20) 26 cussion points. 27 We know from archeological data that in the upper Paleolithic period Ana- 28 tolia was settled by populations with Aurignacian culture (Kuhn 2002). Recent 29 genetic studies (Cinniog¢lu et al. 2004; Olivieri et al. 2006) based on the anal- 30 ysis of mtDNA (haplogroup M1 and U6) and the Y chromosome (R1b3-M269 31 lineage) suggest, in agreement with paleoenvironmental evidence (van Andel and 32 Tzedakis 1996), that around 40,000–45,000 years ago, populations with Aurigna- 33 cian culture may have spread by migration from the Levant and southwest Asia 34 to Anatolia and further into Europe (Bar-Yosef 2002). With the exception of these 35 scarce molecular data, almost nothing is known about the biological features of 36 these early Paleolithic Anatolian foragers. Nevertheless, considering the important 37 demographic processes and biological changes undergone by human populations 38 as a result of later and major events (e.g., the Neolithic transition), we believe that 39 the causes of the observed affinity patterns have to be determined from these later 40 periods. 41 From the Mesolithic to the early Neolithic period different lines of evi- 42 dence support an out-of-Africa Mesolithic migration to the Levant by northeast- 43 ern African groups that had biological affinities with sub-Saharan populations. 44 From a genetic point of view, several recent genetic studies have shown that sub-

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1 Saharan genetic lineages (affiliated with the Y-chromosome PN2 clade; Underhill 2 et al. 2001) have spread through Egypt into the Near East, the Mediterranean area, 3 and, for some lineages, as far north as Turkey (E3b-M35 Y lineage; Cinniog¢lu 4 et al. 2004; Luis et al. 2004), probably during several dispersal episodes since the 5 Mesolithic (Cinniog¢lu et al. 2004; King et al. 2008; Lucotte and Mercier 2003; 6 Luis et al. 2004; Quintana-Murci et al. 1999; Semino et al. 2004; Underhill et al. 7 2001). This finding is in agreement with morphological data that suggest that pop- 8 ulations with sub-Saharan morphological elements were present in northeastern 9 Africa, from the Paleolithic to at least the early Holocene, and diffused northward 10 to the Levant and Anatolia beginning in the Mesolithic. Indeed, the rare and incom- 11 plete Paleolithic to early Neolithic skeletal specimens found in Egypt—such as 12 the 33,000-year-old Nazlet Khater specimen (Pinhasi and Semal 2000), the Wadi 13 Kubbaniya skeleton from the late Paleolithic site in the upper Nile valley (Wendorf 14 et al. 1986), the Qarunian (Faiyum) early Neolithic crania (Henneberg et al. 1989; [555], (21) 15 Midant-Reynes 2000), and the Nabta specimen from the Neolithic Nabta Playa 16 site in the western desert of Egypt (Henneberg et al. 1980)—show, with regard to 17 the great African biological diversity, similarities with some of the sub-Saharan Lines: 377 to 379 18 middle Paleolithic and modern sub-Saharan specimens. This affinity pattern be- ——— 19 tween ancient Egyptians and sub-Saharans has also been noticed by several other 0.0pt PgVar 20 investigators (Angel 1972; Berry and Berry 1967, 1972; Keita 1995) and has been ——— 21 recently reinforced by the study of Brace et al. (2005), which clearly shows that Normal Page 22 the cranial morphology of prehistoric and recent northeast African populations is PgEnds: TEX 23 linked to sub-Saharan populations (Niger-Congo populations). These results sup- 24 port the hypothesis that some of the Paleolithic–early Holocene populations from 25 northeast Africa were probably descendents of sub-Saharan ancestral populations. [555], (21) 26 A late Pleistocene–early Holocene northward migration (from Africa to the 27 Levant and to Anatolia) of these populations has been hypothesized from skele- 28 tal data (Angel 1972, 1973; Brace et al. 2005) and from archeological data, as 29 indicated by the probable Nile valley origin of the “Mesolithic” (epi-Paleolithic) 30 Mushabi culture found in the Levant (Bar Yosef 1987). This migration finds some 31 support in the presence in Mediterranean populations (Sicily, Greece, southern 32 Turkey, etc.; Patrinos et al. 2001; Schiliro et al. 1990) of the Benin sickle cell 33 haplotype. This haplotype originated in West Africa and is probably associated 34 with the spread of malaria to southern Europe through an eastern Mediterranean 35 route (Salares et al. 2004) following the expansion of both human and mosquito 36 populations brought about by the advent of the Neolithic transition (Hume et al. 37 2003; Joy et al. 2003; Rich et al. 1998). This northward migration of northeastern 38 African populations carrying sub-Saharan biological elements is concordant with 39 the morphological homogeneity of the Natufian populations (Bocquentin 2003), 40 which present morphological affinity with sub-Saharan populations (Angel 1972; 41 Brace et al. 2005). In addition, the Neolithic revolution was assumed to arise 42 in the late Pleistocene Natufians and subsequently spread into Anatolia and Eu- 43 rope (Bar-Yosef 2002), and the first Anatolian farmers, Neolithic to Bronze Age 44 Mediterraneans and to some degree other Neolithic–Bronze Age Europeans, show

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1 morphological affinities with the Natufians (and indirectly with sub-Saharan pop- 2 ulations; Angel 1972; Brace et al. 2005), in concordance with a process of demic 3 diffusion accompanying the extension of the Neolithic revolution (Cavalli-Sforza 4 et al. 1994). 5 Following the numerous interactions among eastern Mediterranean and Lev- 6 antine populations and regions, caused by the introduction of agriculture from the 7 Levant into Anatolia and southeastern Europe (Bar-Yosef 2002; Keita and Boyce 8 2005; King et al. 2008), there was, beginning in the Bronze Age, a period of in- 9 creasing interactions in the eastern Mediterranean, mainly during the Greek, Ro- 10 man, and Islamic periods. These interactions resulted in the development of trad- 11 ing networks, military campaigns, and settler colonization (Cruciani et al. 2007; 12 Edwards et al. 2000; Keita and Boyce 2005; King et al. 2008; Lucotte and Mercier 13 2003; Sahoglu 2005; Waelkens et al. 2006). Major changes took place during this 14 period, which may have accentuated or diluted the sub-Saharan components of the [556], (22) 15 earlier Anatolian populations. The second option seems more likely, because even 16 though the population from the Sagalassos territory was interacting with northeast- 17 ern African and Levantine populations [trade relationships with Egypt (Arndt et al. Lines: 379 to 385 18 2003), involvement of thousands of mercenaries from Pisidia (Sagalassos region) ——— 19 in the war around 300 b.c. between the Ptolemaic kingdom (centered on Egypt) 0.0pt PgVar 20 and the Seleucid kingdom (Syria/Mesopotamia/Anatolia), etc.], the major cultural ——— 21 and population interactions involving the Anatolian populations since the Bronze Normal Page 22 Age occurred with the Mediterranean populations from southeastern Europe, as PgEnds: TEX 23 suggested from historical (cf. historical context) and genetic data (Berkman et al. 24 2008; Cinniog¢lu et al. 2004; Di Benedetto et al. 2001; Tambets et al. 2000). 25 Consequently, one may hypothesize as the most parsimonious explanation [556], (22) 26 that sub-Saharan biological elements were introduced into the Anatolian popula- 27 tions after the Neolithic spread and have been preserved since this time, at least 28 until the 11th –13th century a.d., in the population living in the Sagalassos ter- 29 ritory of southwestern Anatolia. This scenario implies that the affinity between 30 Sagalassos and the two sub-Saharan populations (Gabon and Somalia) is more 31 likely due to the sharing of a common ancestor and that the major changes and 32 increasing interactions in the eastern Mediterranean beginning in the Bronze Age 33 did not erase some of the sub-Saharan elements carried by Anatolian populations, 34 as shown by genetic data (e.g., Cinniog¢lu et al. 2004; Luis et al. 2004) and the 35 morphological features of our southwestern Anatolian sample. 36 Northern and central European populations from Germany and Scandinavia 37 are also among the closest to Sagalassos (MMDst = 1.02 and 0.7, respectively; 38 see Table 2 and Figures 2 and 3). The Scandinavian sample shows an affinity to 39 the whole set of populations from the northeastern Mediterranean, whereas the 40 German sample does not show such a pattern and has a strong affinity only to 41 the Sagalassos population. As explained previously, convergent adaptation to lo- 42 cal environment is unlikely to be the main cause of this affinity pattern. Direct 43 contact between Scandinavia and Germany and southwestern Anatolia can also 44 be disregarded because there is no clear evidence known to support this, with the

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1 exception of the Viking raid to Constantinople at the end of the 9th century a.d., 2 which did not result in any kind of settlement process. If the observed affinity 3 pattern seems at first relatively difficult to explain, we should acknowledge that it 4 may be in agreement with the geographic map for the second principal component 5 from the analysis by Cavalli-Sforza et al. (1994) regarding European gene frequen- 6 cies. Indeed, this map shows that populations from western Turkey, Greece, central 7 Germany, and even Scandinavia belong to the same biological distance gradient 8 (cline). The interpretation of this patterning is still the focus of considerable de- 9 bate, and there is no general answer to relate this pattern with a specific historical 10 process or episode (Renfrew 2000). 11 However, previous attempts to explain this affinity pattern have claimed that 12 no archeologically or historically documented process after the Neolithic seems 13 to have had the potential to cause such a clinal biological pattern (Barbujani and 14 Chikhi 2000; Renfrew 2000; Richards et al. 2000). This implies that the close re- [557], (23) 15 lationship detected between the northeastern Mediterranean populations (Greek, 16 Cypriot/Turkish, and Sagalassian) and German and Scandinavian samples might 17 date back to the major population movements that took place during the Paleolithic Lines: 385 to 389 18 colonization of Europe and/or during the demic diffusion of Neolithic farmers ——— 19 (Renfrew and Boyle 2000), but without differentiating between the main demo- 0.0pt PgVar 20 graphic events potentially involved: the upper Paleolithic colonization of Europe ——— 21 (45,000 years ago) (Olivieri et al. 2006), the post–Last Glacial Maximum expan- Normal Page 22 sion (about 20,000 years ago) (Taberlet et al. 1998; Hewitt 2000), the Younger PgEnds: TEX 23 Dryas–Holocene reexpansion from the Anatolian glacial refuge (about 12,000 24 years ago), and the population growth associated with the introduction of agri- 25 cultural practices (about 8,000 years ago) (Ammerman and Cavalli-Sforza 1984). [557], (23) 26 If this scenario is correct, we could hypothesize that northern and central Eu- 27 ropean populations and the population living in Sagalassos have affinities through 28 cranial discrete traits that may have been shaped by the same demographic pro- 29 cess, dating back to the Paleolithic/Neolithic population movements, and that these 30 populations have undergone relatively few changes for the morphological mark- 31 ers analyzed in this study since this major demographic event, which resulted in 32 their patristic similarity. However, different lines of evidence from archeological 33 and historical records and recent genetic studies suggest that subsequent major 34 demographic events occurring in central and southeastern Europe and Anatolia 35 might have had the potential maybe not to cause the affinity pattern observed 36 but probably to accentuate a preexisting pattern. Indeed, recent genetic studies 37 (e.g., Cinniog¢lu et al. 2004; Cruciani et al. 2007) refined previous evolutionary 38 hypotheses and showed that the distribution of some genetic variations (E-V13, 39 J-M12, and M170 Y-chromosome lineages) previously attributed to early popu- 40 lation movements (e.g., Neolithic) into Europe may in fact have resulted from 41 demographic events and population expansion initiated in the Bronze Age. These 42 studies suggest that some genetic lineages underwent a demographic expansion 43 4,000 or 5,000 years ago from the Balkans toward north-central Europe and into 44 western Anatolia (Cinniog¢lu et al. 2004; Cruciani et al. 2007; Semino et al. 2004),

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1 in agreement with the beginning of the Balkans Bronze Age, and underwent sub- 2 sequent demographic expansion in the neighboring region, as suggested by arche- 3 ological and genetic data (Coleman 2000; Cruciani et al. 2007; Kristiansen 1998). 4 This expansion probably followed the major route connecting west Anatolia to 5 the southern Balkans and to north-central Europe used for cultural, material, and 6 gene exchanges to and from northern and central Europe since the spread of the 7 Neolithic (Kristiansen 1998; Mellaart 1968; Tringham 2000). 8 In this context it is likely that Bronze Age events may have facilitated the 9 southward diffusion of populations carrying northern and central European bio- 10 logical elements and may have contributed to some degree of admixture between 11 northern and central Europeans and Anatolians, and on a larger scale, between 12 northeastern Mediterraneans and Anatolians. Even if we do not know which pop- 13 ulations were involved, historical and archeological data suggest, for instance, the 14 2nd millennium b.c. Minoan and later the Mycenaean occupation of the Anatolian [558], (24) 15 coast, the arrival in Anatolia in the early 1st millennium b.c. of the Phrygians com- 16 ing from Thrace, and later the arrival of settlers from Macedonia in Pisidia and in 17 the Sagalassos territory (under Seleucid rule; Waelkens 2004). The coming of the Lines: 389 to 397 18 Dorians from northern Greece and central Europe (the Dorians are claimed to be ——— 19 one of the main groups at the origin of the ancient Greeks) may have also brought 0.0pt PgVar 20 northern and central European biological elements into southern populations. In- ——— 21 deed, the Dorians may have migrated southward to the Peloponnese, across the Normal Page 22 southern Aegean and Crete, and later reached Asia Minor. Nevertheless, the im- PgEnds: TEX 23 pact of the Dorians in the eastern Mediterranean population history is still debated, 24 as the historical, linguistic, and archeological data do not agree (Hall 2006). 25 Consequently, we can postulate that the similarity in cranial discrete traits [558], (24) 26 between Sagalassos and northern and central European populations may have re- 27 sulted from an early (i.e., Paleolithic or Neolithic) shared common ancestor and 28 may have been reinforced more recently through gene flow between these regions 29 or populations. 30 In conclusion, the results of this study suggest that the affinity pattern based 31 on cranial discrete traits of the Byzantine population from Sagalassos (southwest- 32 ern Turkey) can best be interpreted as reflecting multiple demographic events that 33 occurred over the centuries in this region, in part because of its strategic geographic 34 position as a crossroads between Africa and Eurasia. Indeed, our results support 35 the idea that the cranial discrete traits of the 11th–13th-century Sagalassos pop- 36 ulation have retained the traces of major migratory events that have affected the 37 population living in the Sagalassos territory over the last millennia. This implies 38 that the population settled in this area has been relatively continuous and stable, in 39 agreement with archeological data (continuous human occupation of the Sagalas- 40 sos territory since the 12th millennium b.p.) and that cranial discrete trait affinity 41 patterns are best understood within a large interpretive framework, which can then 42 reveal the different strata of the population biological history. 43 Nevertheless, these preliminary results are inevitably constrained by the 44 sample size and other factors, which did not allow us to assess, for instance,

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1 the biological impact of some populations historically involved in the Anatolian 2 ethnogeny (because of the lack of available data from eastern Mediterranean pop- 3 ulations). Also, with regard to our historical overview, we were not able to in- 4 vestigate diachronic variation of the Sagalassian morphology. Only through fur- 5 ther archeological surveys and biological studies from northeastern Mediterranean 6 populations will a better understanding of the Sagalassos population history (and 7 on a larger scale, the regional population history) emerge to validate or invalidate 8 the hypotheses we put forward here. Future excavations of earlier Sagalassos burial 9 sites (8th–5th century b.c.) and the ongoing genetic analysis of human remains 10 belonging to the Sagalassos population should illuminate some of these issues. 11 12 Acknowledgments This research was supported by the Belgian Programme on Inter Uni- 13 versity Poles of Attraction (IUAP V and VI) initiated by the Belgian State, Prime Minister’s 14 Office, Science Policy Programming. We also present the results of a Concerted Action of [559], (25) 15 the Flemish Government (GOA02 and GOA07) and of a project of the Fund for Scientific 16 Research-Flanders (Belgium) (FWO). 17 Lines: 397 to 434 18 Received 11 January 2008; revision received 26 March 2008. ——— 19 -2.83101pt PgVar 20 ——— 21 Literature Cited Normal Page 22 Ammerman, A., and L. L. Cavalli-Sforza. 1984. The Neolithic Transition and Genetics of Populations PgEnds: TEX 23 in Europe. Princeton, NJ: Princeton University Press. 24 Angel, L. 1972. Biological relationships of Egyptian and eastern Mediterranean populations during 25 predynastic times. J. Hum. Evol. 1:307–313. [559], (25) 26 Angel, L. 1973. Early Neolithic people of Nea Nikomedeia. In Fundamenta,v.1,Die Anfänge Ne- olithikums vom Orient bis Nordeuropa 3, I. Schwidetsky, ed. Monographien zur Urgeschichte, 27 ser. B. Cologne: Institut für Ur- und Frühgeschichte, Universität zu Köln, 103–112. 28 Armstrong, J. S. 2007. Significance tests harm progress in forecasting. Int. J. Forecasting 23:321–327. 29 Arndt, A., W. Van Neer, B. Hellemans et al. 2003. Roman trade relationships at Sagalassos (Turkey) 30 elucidated by ancient DNA of fish remains. J. Archaeol. Sci. 30:1095–1105. 31 Barbujani, C., and L. Chikhi. 2000. Genetic population structure of Europeans inferred from nuclear and mitochondrial DNA polymorphisms. In Archaeogenetic: DNA and the Population Pre- 32 history of Europe, C. Renfrew and K. Boyle, eds. Cambridge, U.K.: McDonald Institute for 33 Archaeological Research, 119–129. 34 Bar-Yosef, O. 1987. Pleistocene connexions between Africa and southwest Asia. Afr. Archaeol. Rev. 35 5:29–30. 36 Bar-Yosef, O. 1998. The Natufian culture in the Levant: Threshold to the origins of agriculture. Evol. Anthropol. 6:159–177. 37 Bar-Yosef, O. 2002. The Upper Paleolithic revolution. Annu. Rev. Anthropol. 31:363–393. 38 Bellwood, R., and C. Renfrew. 2002. Examining the Farming/Language Dispersal Hypothesis. Cam- 39 bridge, U.K.: McDonald Institute for Archaeological Research. 40 Berkman C. C., H. Dinc, C. Sekeryapan et al. 2008. Alu insertion polymorphisms and an assessment 41 of the genetic contribution of Central Asia to Anatolia with respect to the Balkans. Am. J. Phys. Anthropol. 136:11–18. 42 Berry, A. C., and R. J. Berry. 1967. Epigenetic variation in the human cranium. J. Anat. 101:361–379. 43 Berry, A. C., and R. J. Berry. 1972. Origins and relationships of the ancient Egyptians. J. Hum. Evol. 44 1:199–208.

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