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Post by Admin on Mar 10, 2020 21:41:39 GMT
Ancient human mitochondrial genomes from Bronze Age Bulgaria: new insights into the genetic history of Thracians Alessandra Modi, Desislava Nesheva, Stefania Sarno, Stefania Vai, Sena Karachanak-Yankova, Donata Luiselli, Elena Pilli, Martina Lari, Chiara Vergata, Yordan Yordanov, Diana Dimitrova, Petar Kalcev, Rada Staneva, Olga Antonova, Savina Hadjidekova, Angel Galabov, Draga Toncheva & David Caramelli Scientific Reports volume 9, Article number: 5412 (2019) Abstract One of the best documented Indo-European civilizations that inhabited Bulgaria is the Thracians, who lasted for more than five millennia and whose origin and relationships with other past and present-day populations are debated among researchers. Here we report 25 new complete mitochondrial genomes of ancient individuals coming from three necropolises located in different regions of Bulgaria – Shekerdja mogila, Gabrova mogila and Bereketska mogila – dated to II-III millennium BC. The identified mtDNA haplogroup composition reflects the mitochondrial variability of Western Eurasia. In particular, within the ancient Eurasian genetic landscape, Thracians locate in an intermediate position between Early Neolithic farmers and Late Neolithic-Bronze Age steppe pastoralists, supporting the scenario that the Balkan region has been a link between Eastern Europe and the Mediterranean since the prehistoric time. Spatial Principal Component Analysis (sPCA) performed on Thracian and modern mtDNA sequences, confirms the pattern highlighted on ancient populations, overall indicating that the maternal gene pool of Thracians reflects their central geographical position at the gateway of Europe. Introduction Bulgaria is situated in the eastern part of the Balkan Peninsula, at the connection point between Southeastern Europe, Eurasian steppe, Anatolia and the Aegean islands. The presence of modern humans in this region is attested starting from 40 kya by the Paleolithic series at Bacho Kiro and Temnata Dupka Caves1,2,3. Some archaeological sites associated with early farmers, as well as the earliest evidence of copper metallurgy in Europe, indicates that this area played a significant role both in the Neolithic and in the Metal Ages4. One of the best documented Indo-European civilizations that inhabited Bulgaria consists in the Thracians, whose cultural legacy is still evident in the modern society. Different theories have been historically proposed about the origin of the Thracians. Today it is assumed that the Thracian culture emerged and formed in the early Bronze Age5,6,7, a period characterized by strong cultural changes and movements of people westward from the Steppe8. During the 5th and 4th millennium BCE, the inhabitants of the eastern region of Balkans were organized in different groups of indigenous people that, over time, were named under the single ethnonym of “Thracians”9,10,11. According to historical and archaeological sources, the Thracian culture flourished during the 2nd and 3rd millennia BCE12,13. The rich cultural and historical heritage, represented by fortresses and necropolises, as well as by the world-famous Panagyurishte, Valchitran, Lukovit and Rogozen treasures, dates back to this period. In the later periods, several populations (Greeks, Macedonians, Slavs and proto-Bulgarians) arrived in the Balkans, reaching the lands occupied by Thracians and mixing with them, thus influencing their cultural and biological identity11. Figure 1 Genetic analyses on both autosomal variations14 and uniparental genetic markers15,16,17 of present-day Bulgarians, locate them between Eastern European and Mediterranean populations, with a particular affinity to the neighboring groups from Greece and the Balkans16. In addition, the Bulgarian maternal genetic pool particularly suggests a major Western Eurasian origin, tracing their ancestry to lineages that witness a complex genetic structure of the region today and reflect different peopling and admixture events from the Upper Paleolithic to the onset of the Neolithic and Post-Neolithic in Europe15. Recent genome-wide ancient DNA (aDNA) based studies on Southeastern Europe, have shown that Neolithic population from present-day Bulgaria was closely related with the northwestern-Anatolian-Neolithic ancestry that signals the spread of Early farmers across Europe, except for the individuals lived in the mid-sixth millennium BC in Malak Preslavets, who revealed a significantly higher level of hunter-gatherer-related ancestry than the other Balkan Neolithic individuals18. Starting from the early 3rd millennium BCE, migrations from the adjacent Pontic-Caspian and Eurasian steppe also played an important role in the transformation of the European genetic landscape, and the contribution of Steppe ancestry to Southeastern European populations increased particularly during the Bronze Age18. Although our understanding of the population and cultural dynamics occurred in the (pre-) history of Balkan Peninsula is starting to be increasingly elucidated, the genetic details on the local civilization remain unknown and this information is only partly available for the ancient (proto-) Bulgarian people18,19. We now have the opportunity to investigate the genetic structure of the Thracians, an ancient people that lasted for more than five millennia and whose origin and relationships with other past and present-day populations are still debated among researchers. To investigate the genetic structure and population history of this ancient civilization, we analyzed 25 complete mitochondrial genomes from three Thracian necropolises (Fig. 1 and Table 1) along with modern and ancient European data. The characterization of the Thracian mitochondrial DNA (mtDNA) variability may have important implications for understanding the dynamics of interaction between Eastern Europe and the Mediterranean, and will also contribute to better clarify the genetic evolution of European populations and the origin of contemporary Bulgarian gene pool.
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Post by Admin on Mar 11, 2020 2:22:16 GMT
Results We successfully reconstructed complete or almost entire mitochondrial genomes for 26 individuals, 3 from Shekerdja mogila, 1 from Gabrova mogila and 22 from Bereketska mogila (Table 2). All the resulted sequences reach the standard quality requested to guaranty the reliability of the NGS data; CtoT patterns range between 20% to 46%, average fragment size vary from 44.4 base pair (bp) to 67.4 bp and no significant levels of present-day human contamination were detected (Table 2). Only one sample, BM-51, showed a high level of contamination and was not considered in the following statistical analyses. The direct radiocarbon dating performed on the samples BM 44, SM 8.1 and GM 30.3 placed the remains at II-III millennium BC (CEDAD, Centro di DAtazione e Diagnostica, Univerità del Salento, Italy) (Table 1), that corresponds to the age estimated according to the archaeological record. The mtDNA sequences obtained were assigned to 21 different haplogroups, representative of the mitochondrial variability of Western Eurasia (Table 2 and Supplementary Table S1). Phylogenetic links between haplotypes of the Thracian samples and comparison ancient data are shown in the Median Joining Network (Fig. 2). Most of the Thracian individuals belong to sub-lineages of the macro-haplogroup H, which accounts for an overall frequency of 33%. This is the most frequent mitochondrial lineage in present-day Europe, representing over 40% of the total mtDNA variability20. Its frequency observed in the Thracians samples is almost similar to the frequency in contemporary European population. Two individuals belong to haplogroup HV, an ancient European lineage likely originating in the Mediterranean region during the Last Glacial Maximum (LGM)21. In ancient samples, HV has been identified in one Mesolithic specimen from Sicily22 and in early Neolithic remains from Spain23, Germany8 and Russia18,24; Mathieson et al.18 reported a HV haplotype in one sample from Serbia dating from 5800 BCE. Moreover, haplogroup HV was observed in Copper Age specimens from Scotland, Hungary and Germany25 and in Hungarian and Israeli samples from the Chalcolithic period26,27 Figure 2 We found four individuals belonging to haplogroup K1c (GM-30.3, BM-51A, BM-58A and BM-68). All the haplotypes contain the expected K1c defining variants with the following private polymorphisms: GM-30.3, 309.1T, 310C, 7441T and 16519C; BM-51A, 16519C; BM-58A, 310C, 513.1CA and 16519C; BM-68, 5297T and 16519C. Nowadays the highest observed European frequency of the lineage K is in Bulgaria (13.3%)28 and K1c is particularly common in Slavic-speaking countries. In ancient populations, the haplogroup K1c has been identified in six hunter-gatherers dated before the arrival of farming (one in Romania, three in Serbia18 and two in Greece29), in two Bronze-Age individuals from Hungary and Bulgaria18,30,31 and in two Central-Europe farmers associated with the Bell-Beaker culture25,32,33. The phylogenetic network analysis (Fig. 2) reveals that the detected K1c haplotypes in Thracians are closely related to hunter-gatherers from Iron Gates and Bronze Age individuals from Bulgaria and Hungary. Three samples belong to haplogroup J1c (SM-4, BM-31 and BM-61). The SM-4 individual shows three personal transitions previously identified at positions 199C, 8730G and 13928A, and a private mutation at 13686G. The haplotypes of samples BM-31 and BM-61 fall within the sub-haplogropus J1c9 and J1c6, respectively. Currently, J1c, which dates to ∼16 ka ago, is found mainly in Europe, especially in Central Europe, Balkans and Ukraine, where it encompasses almost 80% of total J1 lineages. Pala et al.34 suggested that during the LGM, haplogroup J sub-lineages arose in the Near Eastern refugia and recolonized Europe following the end of the last glaciation. In particular, J1c is not yet found in any hunter-gatherers, and the oldest individuals belonging to this lineage were found in Iran35 and in Anatolia30 dating to 8000-7700 BCE. It is possible that J1c arrived in Thracia from Anatolia during the early stages of the Neolithic expansion. The expansion of farmers played an important role also in the diffusion of haplogroup T, which has been found in three Thracian samples with the T2b (BM-15 and BM-59A) and T2e (BM-40) sub-lineages. Pala et al.34 particularly suggested that these lineages entered Europe from Anatolia in the Late Glacial period, and have been later diffused around Europe by Neolithic agriculturalists after intermingling with the inhabitants of Southeast Europe. Overall, while haplogroups H, K, J and T arose throughout the Neolithic increasing frequencies in different later communities and present-day European populations, the haplogroup U sub-lineages including U2, U4, U5 and U8 instead mark the genetic pool of European pre-LGM hunter-gatherers36,37,38. The mtDNA genetic relationships between Thracians and the other ancient Eurasian populations (Supplementary Table S2) were directly explored through a correspondence analysis (COA, Fig. 3). The first component, which accounted for 28.3% of the total variance, clearly separates all hunter-gatherers from the rest of Neolithic, Bronze Age and Iron-Age population groups. Along the second component (10,6% of variance), the ancient populations appear instead distributed along a cline of genetic variation which extends from the Early Neolithic farmers of Southern Europe and Anatolia to the Late Neolithic/Bronze Age Europeans and Steppe pastoralists, in accordance with the genomic structure of ancient Europe29,30,32,33. From an autosomal genetic perspective, besides showing the clear discontinuity of Paleolithic hunter-gatherers, recent genome-wide aDNA studies, have indeed outlined two opposite genetic components contributing to the European genetic ancestry: i.e. the ancestry of the Early European farmers related to Anatolian farmers and pre-farming Levant populations and, on the other side, the so-called Steppe ancestry eventually spread into Europe and Asia during the Bronze Age migrations of Yamnaya herders. In this scenario, the mtDNA genetic composition of analyzed Thracian population located them in the middle of this cline, clustering closely to the Peloponnese-Neolithic individuals (Peloponnese_N) and the Chalcolithic and Bronze Age populations of the Balkans (Balkans_Chalcolithic, Balkans_BA). This finding seems to support a mitochondrial genetic profile of the Thracians that reflects their geographical position at the gateway of Europe. In a more general perspective, Thracians show a mtDNA genetic composition that is thus intermediate between the western Eurasian and the Mediterranean populations, documenting a prolonged interaction between people of these regions during the Bronze Age. On the other hand, the relatively higher distance with the Bronze Age populations from the Steppe (Steppe_EMBA and Steppe_MLBA), may support the hypothesis that the Thracians largely derived from local people9,10,11 with only a low percentage of the gene flow from the Steppe, at least during the early stages of their cultural development. However, in order to better explore this hypothesis, it is worth emphasizing that the perspective offered here by the analysis of mitochondrial genomes should be integrated by the possibility of testing the results obtained with Y-chromosome and autosomal genome-wide data. At this respect, several studies have indeed pointed out the sex-biased nature of the recent demographic changes and expansions in Eurasia39,40,41,42,43, thus suggesting possible sex-specific patterns of migration. Figure 3 In addition to a temporal frame, in order to explore the spatial pattern of mtDNA genetic variability, the genetic composition of past Thracian population was compared also with that of present-day human groups by means of a spatial Principal Component Analysis (sPCA, Fig. 4). Along the first component (sPC1) the ancient Thracians are closely related with Central-East European populations, while along the second component (sPC2) our samples show higher resemblance with present-day Mediterranean groups. Despite the general lack of statistical support to a clear-cut genetic structure (Gtest: obs = 0.196, P-value = 0.182), as expected due to the well-known higher genetic homogeneity of the mtDNA variability, this pattern reflects the one highlighted by COA analysis on ancient populations. Overall, the mitochondrial genetic structure observed in our sample seems to be mainly a consequence of demographic processes between two macro-areas: West Eurasia and the Mediterranean. This is in agreement with previous studies on modern samples14,15,16 that identify features of both Eastern Europe and Mediterranean area in Bulgarian population. Figure 4
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Post by Admin on Mar 11, 2020 5:23:56 GMT
Discussion In the present study, we reconstructed and analyzed complete mitochondrial genomes from 25 Bronze Age individuals sampled in three Bulgarian necropolises. According to the archaeological records, these cemeteries are associated to the Thracians culture and the chronology, attributed by funerary context, was confirmed by three direct radiocarbon dating placing the remains at II-III millennium BC. These data were used to explore, for the first time, the genetic structure of this ancient population.
We found that the Thracian maternal gene pool is represented essentially by Western Eurasian haplogroups, as expected given the well-known overall mtDNA genetic similarity among all European populations. However, when we compared the complete mitochondrial sequences of Thracians to that of ancient and contemporary Eurasian populations, we observe that their genetic profile reflects their nexus geographical position between east and west.
Several studies demonstrated that Balkan Peninsula has been in different times a crossroad for people moving from and to Europe and beyond16,44. While previous analyses of modern populations demonstrated the impact of such migrations on the genetic makeup of present-day Bulgarians14,15,16, scarce information were available for the ancient (proto-) Bulgarian maternal gene pool and were mainly limited to HVS1 data from the medieval period19. In this study, we provide, for the first time, genetic details of an ancient population, which is particularly relevant from both a chronological and a geographical point of view. In accordance with their geographical location, Thracians show a genetic composition clearly intermediate between East Europe and Mediterranean, that suggests multiple admixture events and population movements occurred across what is now the modern day Bulgaria. Albeit limited to DNA transmitted along the female lines of descent, our genetic data on ancient Thracians provide a direct evidence of how the Balkan region has been a link between East and West Europe since the prehistoric time, and particularly during the Neolithic and post-Neolithic events. In this perspective, future studies will certainly benefit from the analysis of nuclear genome (Y-chromosome and autosomal genetic variation) in order to integrate the observed mtDNA genetic patterns within a more comprehensive overview and for testing the possibility of different sex-biased migrations in the area.
Overall, the ancient mtDNA data presented in this study integrate the existing database and has important implication for understanding the origins of the peopling in this part of Europe and for enlarging the knowledge on the ancient Bronze Age civilizations. How and to what extent ancient Thracian people has contributed to the present-day Bulgarian gene pool remain largely unknown due to the lack of large mitogenomes from contemporary populations from the area, necessary for a phylogenetically and demographically informative comparison.
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Post by Admin on Mar 17, 2020 20:50:43 GMT
Y-Chromosome Analysis in Individuals Bearing the Basarab Name of the First Dynasty of Wallachian Kings Begoña Martinez-Cruz, Mihai Ioana, [...], and the Genographic Consortium Abstract Vlad III The Impaler, also known as Dracula, descended from the dynasty of Basarab, the first rulers of independent Wallachia, in present Romania. Whether this dynasty is of Cuman (an admixed Turkic people that reached Wallachia from the East in the 11th century) or of local Romanian (Vlach) origin is debated among historians. Earlier studies have demonstrated the value of investigating the Y chromosome of men bearing a historical name, in order to identify their genetic origin. We sampled 29 Romanian men carrying the surname Basarab, in addition to four Romanian populations (from counties Dolj, N = 38; Mehedinti, N = 11; Cluj, N = 50; and Brasov, N = 50), and compared the data with the surrounding populations. We typed 131 SNPs and 19 STRs in the non-recombinant part of the Y-chromosome in all the individuals. We computed a PCA to situate the Basarab individuals in the context of Romania and its neighboring populations. Different Y-chromosome haplogroups were found within the individuals bearing the Basarab name. All haplogroups are common in Romania and other Central and Eastern European populations. In a PCA, the Basarab group clusters within other Romanian populations. We found several clusters of Basarab individuals having a common ancestor within the period of the last 600 years. The diversity of haplogroups found shows that not all individuals carrying the surname Basarab can be direct biological descendants of the Basarab dynasty. The absence of Eastern Asian lineages in the Basarab men can be interpreted as a lack of evidence for a Cuman origin of the Basarab dynasty, although it cannot be positively ruled out. It can be therefore concluded that the Basarab dynasty was successful in spreading its name beyond the spread of its genes. Figure 1 Map with the geographical location of the populations sampled. Introduction Vlad III The Impaler, commonly known in the popular literature as Dracula, was a 15th century prince of Wallachia, in current southern Romania. As a ruler, he fiercely resisted the Ottoman expansion. He infamously applied cruel punishments to his enemies and to traitors, including massive impalements that gave rise to his dark legend. His dynasty, the Basarab, took its family name from the first ruler of Wallachia, Basarab I, who rebelled against Charles I of Hungary and in 1330 gained the independence of the country from the Kingdom of Hungary. The dynasty ruled until the assasination of Michael the Brave in 1601 [1]. The name Basarab most probably means father ruler in the Turkic Cuman language [2]. Cumans were a confederation of two different people: the Cuman people that came from the east of the Yangtze River, and the Kipchak people, a Turkic tribal confederation, which occupied a vast territory in the Eurasian steppe, from north of the Aral Sea to the north region of the Black Sea [3]. They expanded into Moldavia, Wallachia and Transylvania by the 11th century, influencing the politics of the region and establishing several royal dynasties, one of which may have been the Basarab of Wallachia [3]. Otherwise, contemporaries identified Basarab I as a local Romanian or Vlach (the local Romanic-speaking population), as Charles of Hungary referred to him as “Basarab our unfaithful Vlach” [3]. Whether the dynasty was of Cuman or Romanian/Vlach origin is a subject of intense debate among historians [1], [2], [3], [4]. Patrilineal surnames mirror the inheritance of the non-recombinant part of the Y-chromosome (NRY), making surnames as markers of male ancestry useful to help answering questions on the history and structure of populations in combination with genetic studies. Additionally, the study of the NRY lineage in males with the same surname could shed light on the history of lineages bearing this name. Males with identical patrilineally inherited surname descending from a common male ancestor will carry the same Y-chromosome lineage, and share higher levels of co-ancestry among them than with the rest of individuals carrying the same NRY lineage in the population [5]. However, there are confounding factors including multiple male founders for the same surname, extra-marital paternity, drift, and surname change from one generation to the next [5]. Another limiting factor of using surnames as genetic markers is the time depth of inherited surnames, which is highly variable across countries and populations [5]. The study of present day Romanian Basarab genetics has the potential to answer interesting questions about the history of the dynasty of the famous Romanian prince. Here we present a study on the Y chromosome of 29 individuals carrying the Basarab surname in present Romania and in 484 individuals from four Romanian and three neighbour populations (Bulgaria, Ukraine, and Hungary) by typing 131 SNPs and 17 STR markers in the NRY. Under the hypothesis that carriers of the Basarab name at present could be the direct descendants of the first ruler of Wallachia, we wanted to ask the following questions: are individuals presently bearing the Basarab surname actually the direct descents of Basarab I? If so, which was the Y-chromosome lineage of the dynasty? Are the Basarab individuals of Cuman or Romanian/Vlach origin?
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Post by Admin on Mar 18, 2020 5:36:08 GMT
Results Descriptive statistics for the studied populations (Figure 1) are shown in Table 1, and haplogroups and haplotypes for the Basarab and the other Romanian populations are given in Table S1. Haplogroups and haplotypes for the surrounding populations of Bulgaria, Hungary and Ukraine are given in Table S2. In general, the Basarab show lower levels of haplotype diversity compared to neighboring samples. Haplotype and haplogroup diversity is especially low in Basarab, and this may be due to founder effects within bearers of the same name. Otherwise, haplogroup diversity in the Romanian populations in this study is higher than previously reported in other Romanian populations (0.7828 in Constanta, 0.8048 in Ploiesti, [6]) but this may be due to the lower level of phylogenetic definition in the previous study. PCA based on haplogroup frequencies grouped the Basarab with the southern Romanian populations of Dolj, Brasov and Mehedinti, and the Ukrainian population. The Romanians from Cluj, the Hungarians and the Bulgarians were more scattered in the plot (Figure 2). The Basarab individuals appear just in the middle of Romanian populations, indicating a common genetic background. Figure 2 PCA plot based on haplogroup frequencies in Basarab, four Romanian populations, and the general populations from Bulgaria, Hungary and Ukraine. The Basarab sample clusters into 11 lineages (Table 2), with six main lineages comprising 82.8% of the samples. Some lineages such as J-M241 and E-V13 are over-represented in the Basarab compared to the general Romanians. The age of each cluster was computed with the ρ method, as in [7]. All the lineages in the Basarab show clear geographical clustering but, with one possible exception, none of them reaches the Middle Ages. It is worth noting that these lineages include a few non-Basarab individuals, but they are all from Hungary or Bulgaria. None is an exact haplotype match, and these coincidences may be caused either by sharing a recent common ancestor with the Basarab, or by the homoplastic nature of STR mutation. Three Basarab founding lineages are found in haplogroup E1b1b1a2-V13; if they were considered together, their joint age would be 1740±615 years, well beyond the establishment of the Romanian nobility. Similarly, if the two lineages in haplogroup I2a-P37.2 were pooled, their joint age would be 960±480 years, or 250 years before the actual founding of the Basarab dynasty. Only Basarab individuals were considered in the age estimations. Lineages found in the Basarab sample. Lineage Haplogroup individuals origin age±SD 1 E1b1b1a2-V13 RU226* Sibiu Basarab 150±150 ya RU231* Sibiu Basarab RU234* Sibiu Basarab RU247 Sibiu Basarab 2 E1b1b1a2-V13 RU239* Gorj Basarab 200±200 ya RU240* Gorj Basarab BM024 Bulgarian 3 E1b1b1a2-V13 RU221* Sibiu Basarab 240±120 ya RU228* Sibiu Basarab RU232* Sibiu Basarab RU233* Sibiu Basarab BM072 Bulgarian 4 I2a-P37.2 RU227* Sibiu Basarab 150±150 ya RU229* Sibiu Basarab RU230* Sibiu Basarab BM019 Bulgarian 5 I2a-P37.2 RU246* Caras-Severin Basarab – RU245* Mehedinti Basarab 6 J2b2-M241 RU219* Sibiu Basarab 200±115 ya RU220* Sibiu Basarab RU222* Sibiu Basarab RU223* Sibiu Basarab RU224* Sibiu Basarab RU235* Sibiu Basarab RU236* Sibiu Basarab RU225 Sibiu Basarab RU241 Gorj Basarab 7 G2a-P15 RU237 Bacau Basarab – 8 J1e-P58 RU242 Gorj Basarab – 9 J2a2-M67 RU238 Bacau Basarab 300±300 ya HM045 Hungarian 10 R1a1a-M17 RU243 Ilfov Basarab 600±283 ya HM213 Hungarian HM162 Hungarian 11 R1a1a7-M458 RU244 Ilfov Basarab – (*): individuals within the same lineage carrying the same haplotype.
Discussion The presence of different Y-chromosome lineages among the individuals that currently carry the name Basarab indicates that not all of them could be direct descendents of the dynasty. Extra-pair paternity could explain the existence of highly different male lineages in a dynasty, but only a very high rate could explain the diversity found in the Basarab population studied. Otherwise, descendants of the Craioveşti boyars/noblemen, a family that claimed direct descent from the Basarab House, may have kept the Basarab name, adding diversity to the Y lineage. Indeed, the genetic evidence indicates that Basarab is a polyphyletic name, with multiple male founders that would explain the pattern of diversity. The use of nicknames to distinguish among individuals with the same given name was common in Romania in the past centuries [8]. Although the most common nickname was the patronymical, others designated the place of origin. Later on, these nicknames became family names. Basarab may thus also indicate a demonym for the historic region of Basarabia (roughly the currently independent Moldova and part of southern Ukraine), reversing the etimological pathway, since the region was first named after the House of Basarab [3]. Additionally, the name could have been adopted as a mark of distinction, given its noble origin. The time depth of inherited surnames is highly variable across countries and populations [5]. In Romania, it was not until 1895 (Law on the name, nr. 18/March 1895) that the first law obliging people to have a first name and a surname was passed [9]. However, in the rural areas this regulation was not effectively applied until two or three decades after. Therefore, the expected time depth of inherited surnames in Romania should be around 100–150 years. The time depth estimated for most (although not all) of the common lineages in the Basarab is in agreement with the time of establishment of surnames in Romania, as seen before in other populations [10]. Only one of these lineages within Romania dates back to medieval times. Interestingly, two Hungarian individuals share this haplotype, and it is well known that a major migration of Cumans took place from the actual territory of Romania to Hungary in the 13th century, where they asked protection from the Hungarian kings against the advancing Mongol invasion [4]. Although tempting, it is impossible to clearly link this particular Y haplotype to a Cuman origin. Nonetheless, we cannot rule out the possibility that one of the Y-chromosome lineages found in the Basarab was indeed the lineage carried by the dynasty. Unfortunately, given the results obtained in this study, only the analysis of the remains of Basarab I or any of his known descendants could confirm or not this hypothesis. Although Cumans came from East Asia, other authors have reported that they also showed Caucasoid features [3]. Historians agree that Cumans mingled with the populations they encountered [3]. West Eurasian Y-chromosome haplogroup R1a1 has been found in admixed East Asian populations as early as in the early Bronze Age [11]. The single study on the genetics of Cumans [12] was based on mitochondrial DNA (mtDNA) and showed that just one individual out of 11 in a medieval burial in Hungary did not carry a Western Eurasian but an East Asian haplogroup (haplogroup D). Otherwise, D is also one of the most frequent mtDNA haplogroups in southern Siberia [13]. However, one can speculate that, given the political dominance of the Cuman, asymmetrical admixture would preserve the Eastern lineages more readily in the NRY than in mtDNA. Thus, we could attribute a Cuman origin to a Basarab lineage if it belonged to an East Asian haplogroup, but a European haplogroup could be carried both by the Cumans and by the native Romanians/Vlachs. As shown in the PCA, the haplogroup composition of the Basarab is very similar to that of the general Romanian population, and none of the haplogroups they carry are particular of Central or East Asia. Therefore, our results are consistent both with an ethnic Cuman or a Romanian/Vlach origin. On the other hand, the extensive presence of Western Eurasian haplotypes in both known medieval Cuman burials and in individuals bearing the Basarab name suggests a significant probability that Basarab I may also have been carrying a Western Eurasian haplotype. To the best of our knowledge, this is the first genetic study on the surname of a royal dynasty. We have shown that not all the people in Romania that bear the name Basarab are direct descendants of the dynasty of the first rulers of Wallachia. It seems that the House of Basarab was rather more successful in extending its name than in passing down its genes.
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