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Post by Admin on May 3, 2022 17:39:26 GMT
Nagykörü-Május 1. úti bölcsőde, grave 6 (NM/6): N3a4-Z1936 (xL1034), Y13850 Based on 14 markers, the study sample fully matched to a sample from Mureş County, Romania (Transylvania) in FTDNA’s database (see Fig. 6 mark c), whose SNP tests showed it to belong to the Z1936 + Y13850 + L1034- group. This group is the ancient relative of the Ugric L1034 group, which is found in modern Volga Tatars from the Republic of Tatarstan. The SNP-based age of its common ancestor with L1034 is 4300 ± 800 years, placing the mutations of Y13850 and L1034 around the time of the divergence of the Baltic-Finns (N3a4-B535 branch) from the Ugrians (N3a4-Y13850 branch). This sample shares common paternal ancestors with Ugric-speaking individuals from Western Siberia. The modern Hungarian N3a4-Z1936*(xL1034) sample from Bodrogköz (Pamjav et al. 2017) is only one mutation away from the research sample in the Z1936 network.
Tiszakécske-Ókécske, grave 1 (TÓ/1): N3a4-L1034 This is the only sample from the analyzed Hungarian Conqueror remains that tested positive for the L1034 group. As such, it connects modern Hungarians and ethnic Székelys with the West Siberian Mansi (Fehér et al. 2015). The sample is genetic one step away (STR difference of 1) from a Greek N3a4-L1034 sample.
Nagykőrös-Fekete dűlő, grave 1 (NF/1): J1-M267 (xL620) This sample belongs to the J1 subgroup. The J-L136/J-P58 subgroups (11,000–12,000 years old) are characteristic of Semitic peoples (Arabs and Jews) and the so-called Cohen modal haplotype also belongs here. On the other hand, the Z1828 subgroup (8000 years old) is dominant among the peoples of Dagestan in the Northeast Caucasian Mountains. Based on the results from the STR analysis, the Hungarian Conqueror sample belongs neither to the J-L1936/J-P58 nor to the Z1828 subgroup. Its closest STR-based genetic relatives are members of a rare subgroup only found in Great Britain, with the L620+ FGC6064+ L136- Z1828- mutation. Nevertheless, the Hungarian Conqueror from the Nagykőrös-Fekete dűlő grave is negative for L620 and P58, while it is very distant from the Z1828 Dagestanian subgroup as well.
Nagykőrös-Fekete dűlő, grave 2 (NF/2): R1a-Z93 We compared 20 loci from this sample with over 4200 37-loci haplotypes in FT DNA’s database. We determined that it was 3 genetic steps away from its closest Z93 relatives: 1 South Siberian Khakass, 2 Saudi Arabians, 1 Syrian Arab, 1 German, 1 Pole, 1 Briton, and 1 Iranian (Khorasan Province). In fact, the closest genetic relatives of the study sample belonged to either the Z93* (Z94-) or the Z2125 (common among Iranian and Turkic peoples) group. A Kyrgyz individual from subgroup Z2125 is 1 genetic distance away on 17 markers from the sample published in Underhill et al. (2015). The patrilineal branch of the study sample hails from the Altai-Tian Shan region, of possible Turkic origin.
Karos-Eperjesszög II, grave 61 (KEII/61): R1a-Z93 We compared 15 loci from this sample with over 4200 37-loci haplotypes from FTDNA’s databaseFootnote5 and determined that its closest Z93 relatives were 2 genetic steps away. One was an Arabic individual, but the other’s origins could not be identified. Both relatives belonged to the Z2125 subgroup, which is common among Iranian- and Turkic-speaking peoples (Underhill et al. 2015).
Among the 12 loci from the Z93 samples examined by Underhill et al. (2015), several are 1 genetic step away from the Karos-Eperjesszög II sample. Using DYS393 = 13, the sample has 4 full matches (1 Caucasian Mingrelian (Georgian), 1 Iranian, 1 Tajik, and 1 Kyrgyz), while using DYS393 = 14, it has 5 matches that are 1 genetic step away from the study sample (2 Tajiks, 2 Pashtuns, and 1 Swiss Z93 sample). It is also noteworthy that the sample is 1 genetic step away from a Khazar sample (Klyosov and Faleeva 2017) and that only their DYS385a data differ. Therefore, this sample could be of Turkic/Khazar origin, as supported by the shared historical connection between the Hungarians of the steppe and the Khazars.
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Post by Admin on May 3, 2022 18:30:41 GMT
Karos-Eperjesszög II, graves 16 (KEII/16) and 52 (KEII/52); Karos-Eperjesszög III, grave 11 (KEIII/11): I2a1-L621, CTS10228 The three samples were identical on those loci which were included in the network. One sample was different from the two others on DYS518, but that locus was not included in the network. Regardless of this single step distance, we can consider the three males close relatives. As the matrilineal lineages (mtDNA X2f) of the individuals from grave 52 in Karos II and grave 11 in Karos III are also identical (Neparáczki et al. 2018), they can be considered full siblings, and the grave goods discovered suggest that this genetic lineage belongs to the chief of the Karos-Eperjesszög settlements. The haplotype of the individual from grave 12 in Karos III (Neparáczki et al. 2017) is likewise identical to that of the 3 individuals we examined here, and 11 and was therefore also part of the chief’s family. The I2a1-L621 sample from grave 17 in Karos III (Neparáczki et al. 2017), however, was not a close genetic relative, because it differed on several markers. We looked at 16 loci from 640 I2a-L621 samples in FTDNA’s I2a project database and found that 7 individuals were 2 genetic steps away the Karos samples, of whom 1 was a Hungarian from Kunszentmárton, 2 were Ukrainians, 1 was Lithuanian, 1 was Belarusian, 1 was Russian, and 1 was a German from Poland. Based on SNP analysis, the CTS10228 group is 2200 ± 300 years old. The group’s demographic expansion may have begun in Southeast Poland around that time, as carriers of the oldest subgroup are found there today. The group cannot solely be tied to the Slavs, because the proto-Slavic period was later, around 300–500 CE. Furthermore, the A2512 subgroup is typically Mediterranean (Greek, Jewish diaspora). We compared 15 loci from our data with Rębała et al.’s (2013) samples and found that 3 Poles and 2 Slovaks are 1 genetic step away, while 2 other Poles are 2 genetic steps away. The Karos samples’ STR data are 1 genetic distance on 17 loci in the Balkans to a Bulgarian from Montana and 2 mutation steps to a Bulgarian from Sofia, a Bulgarian from Plovdiv, and a Tuscan Albanian (see Fig. 7). Fig. 7 The network of the I2a1-L621, CTS10228 haplogroup. The haplotype of Karos samples is marked with an arrow. Source of haplotypes: Di Cristofaro et al. (2013); Karachanak et al. (2013); Mielnik-Sikorska et al. (2013); Mirabal et al. (2010); Niederstätter et al. (2012); Pamjav et al. (2017); and Sarno et al. (2016) As all these hits fit into the time period of the Hungarian Conquest, they may be descendants of the tribes. We determined that the Kunszentmárton Hungarian sample belongs to the A815 subgroup. This is interesting, because this subgroup is also found in Moravia, Slovakia, and Ukraine, and it has a specific North Caucasian Karachay subgroup, as well.Footnote6
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Post by Admin on May 3, 2022 19:48:22 GMT
Tiszaeszlár-Bashalom, grave 13 (TBa/13): R1b-L23, Z2106 This sample belongs to either the Indo-European R1b-M269 subgroup’s L23(xM412) haplotype or to the Z2106 subgroup (see Fig. 8), which is commonly referred to as “Eastern European R1b.” It is found extensively among Bashkirs (Burzyansky), Armenians, various Northern Caucasian peoples, Albanians, and Greeks but in a less degree among Iranians, Eastern European Slavs, and Hungarians (Myres et al. 2011). This group dominates the skeletal remains that are considered as proto-Indo-European of the early Bronze Age Pit Grave (Yamna) culture (Allentoft et al. 2015) Fig. 8 The network of the R1b-M269 haplogroup. We placed the haplotype among Myres et al.’s (2011) L23(xM412) samples (n = 203 individuals). The sample from the Tiszaeszlár cemetery is a full genetic match with a modern Hungarian sample from Bodrogköz (Pamjav et al. 2017) on the 17 markers used in the network (marked with arrow). There is a 1–1 mutation step deviation on 2 frequently mutating markers (DYS570 and DYS576) that are not in the network, which is in accord with the observation that the male from Bodrogköz is a direct descendant of either the research individual or the research individual’s close genetic relative. The two samples are 1 genetic step away on 17 loci from a Northeast Caucasian (from Dagestan) Avar sample; this haplogroup may have entered the Hungarian Conqueror genome in the foreground of the Caucasus Rétközberencs-Paromdomb, graves 1 (RP/1) and 2 (RP/2): G2a2-U1, L1266 The genetic markers of the two male individuals from Rétközberencs-Paromdomb graves 1 and 2 are identical, thus, they might be close relatives. The samples’ haplotype may belong either to the G2a2-L1259 group U1(xL13) that had spread with the early agriculturers or to the L1266 subgroup (see Fig. 9), which, according to Rootsi et al. (2012), gave rise to the patrilineal branch of the Northwestern Caucasian peoples (Abkhazians, Abazins, Adyghes/Circassians, and Kabardians). This group also admixed significantly to the gene pool of Northern Caucasian Turkic-speaking peoples (Karachays, Balkars, Kumyks, and Nogais). Fig. 9 Network of the G2a2-U1 haplogroup. There are no complete matches in the networks of either the YHRD database (https://yhrd.org/) or Rootsi et al.’s (2012) STR database (n = 124 individuals) for the analyzed Hungarian Conqueror samples (marked with an arrow). The closest match is 2 genetic steps away. This puts the common ancestor of the Hungarian Conqueror and the 2 Abkhazian, 1 Abazin, 1 Karachay, and 1 West Chinese Uyghur samples in the early Khazar period. Interestingly, despite the Caucasian nature of the group, the founder haplotype matches only with a modern Hungarian and an Estonian sample (https://www.familytreedna.com/public/G-L293/default.aspx?section=yresults), from which the Hungarian Conqueror samples are found 3 mutation steps away. The group could have been incorporated into the Hungarian Conquerors’ gene pool in the foreground of the Caucasus
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Post by Admin on May 3, 2022 21:59:23 GMT
Karos-Eperjesszög I, grave 3 (KEI/3) and Rakamaz-Túróczi part, grave 7 (RT/7): G2a1-P18, L293 These two skeletal remains yielded results on comparatively few loci, thus, a network for them could not be created. Based on the existing STR analyses, they most likely belong to the G2a1-L293 subgroup, which is characteristic of the Northern Caucasian, Iranian-speaking Ossetians. Ten markers from the Rakamaz skeletal remains were examined and yielded 7 matches among the ethnicities from Balanovsky et al.’s (2011) study: 1 Ossetian P18 sample (2 genetic steps away), 4 Ossetians, 1 Dagestanian Avar, and 1 Abkhazian P18 sample (3 genetic steps away). Only seven markers from the Karos skeletal remains could be compared with Balanovsky et al.’s (2011) study, of which the Karos grave 3 sample is a full genetic match with 1 Ossetian, 1 Abkhazian, and 1 Circassian. A further 9 Ossetians, 5 Chechens, 3 Circassians, and 2 Abkhazians are 1 genetic step away. However, due to the low number of comparable markers, these matches do not necessarily suggest a close genetic relationship. None of the G2a1-L293 samples has a known modern Hungarian descendant, nor can they be relatives due to the multiple genetic differences. Genetic distance calculations In our analysis of the Hungarian Conquerors’ ancestry, FST (fixation index) genetic distances and p values were calculated between Y-chromosomal packages of 73 ethnic groups, including modern Hungarian populations and Hungarian Conqueror samples. The modern Hungarian population is represented by the Bodrogköz (HUB) (Pamjav et al. 2017), Csángó (CNG) (Fehér et al. 2015), and Sekler/Székely (SE2) (Csányi et al. 2008) ethnic groups. In the comparative study, the populations were characterized by the frequencies of haplogroups (see Online Resource 5; ESM_5). The Hungarian Conquerors (HUC) are represented by 16 independent paternal lineages. As shown in the median-joining network analyses, the Hungarian Conquerors are particularly heterogeneous and their Y chromosomes originate from several different places, rather than from a single region, so we have separated the HUC samples into two, more homogeneous subgroups: the Northern Pontic (HUP) and the Ural-Altaic (HUA). 2555 out of a total 2628 FST genetic distance calculations had statistically significant values (p < 0.05), while 73 comparisons had a p value above the threshold (see Online Resource 6; ESM_6). The high p values may be explained by nearby populations having such low FST genetic distances so as to be insignificant (e.g., SKW-POL FST = 0.00479; p = 0.18919), or one of the compared populations’ sample sizes being too small (e.g., HUP-ARM FST = 0.07899; p = 0.10811). The highest p values are explained by a combination of the aforementioned reasons (e.g., HUA-UDM FST = 0.01763; p = 0.24324). Some of the FST values are negative, which suggests that the interpopulational genetic distances are smaller than the intrapopulational genetic distances. 15–15 comparisons were insignificant between the modern populations and the two Hungarian Conqueror groups (HUP and HUA). There is no overlap between the HUP and HUA related populations, all 30 are unique (see online Resource 6; ESM_6 and Fig. 10). Fig. 10 A two-dimensional NMDS plot depicting the FST data based on haplogroup frequency. The samples whose FST genetic distances are statistically insignificant to the Ural-Altaic (HUA) and the Northern Pontic (HUP) subgroups are marked by the red and green points, respectively. The modern Hungarian population representing ethnic groups (Bodrogköz (HUB), Csángó (CNG), and Sekler/Székely (SE2)) are marked by blue The FST values between the Ural-Altaic subgroup (HUA) and Karelians (KRL), Estonians (EST), Saami (LAP), Eastern Finns (FIE), Northern Russians (RUN), Lithuanians (LIT), Latvians (LAT), Besermyan (BES), Udmurts (UDM), Komi-Zyrians (KOM), Mari (MRI), Chuvash (CHU), Tatars (TAT), Khanty (KHA), and the Buryats (BRY) are not statistically significant. The FST values between the Pontic Hungarian Conqueror subgroup (HUP) and the Cherkess/Circassians (CHR), Kabards (KAB), Adyghe/Circassians (ADG), Balkars (BLK), Abkhazians (ABH), Ossetians (OSE), Karachays (KAR), Kuban Nogais (NKU), Kumyks (KUM), Georgians (GEO), Armenians (ARM), Sardinians (SRD), Middle-Neolithic Europeans (MDN), Neolithic Hungarians (NEH), and the Neolithic German and Spanish (NGS) samples are not statistically distinguishable. Genetic distances are depicted on an NMDS plot (Fig. 10). HUA and HUP groups are separately placed with distinct neighboring populations. The Ural-Altaic (HUA) subgroup fits within the Finno-Ugric-speaking peoples (Estonians, Karelians, Eastern Finns, Saami, Besermyan, Komi, Udmurts, Khanty, and Mansi) and is close to their neighbors, the Tatars, Chuvash, Buryats, Latvians, Northern Russians, and the Lithuanians. The Northern Pontic (HUP) samples fall far away from the other Hungarian Conqueror subgroup samples and also appear in other ethnic settings. Their closest ethnic groups are predominantly Caucasian (Kabardins, Adyghe/Circassians, Balkars, Cherkess/Circassians, Abkhazians, Georgians, Karachays, Kumyks, and Kuban Nogais). Based on the high frequency of Hg G, the Hungarian, the Middle-Neolithic European, the Neolithic German and Spanish, the Maltese, and the Sardinian groups are also close to each other.
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Post by Admin on May 4, 2022 17:51:16 GMT
Discussion Several studies on the genetic origins of Hungarians have been published within the last decade, but the number of unanswered questions remained high. Furthermore, past studies on skeletons from the Hungarian Conquest Period focused on the maternal lineage, whereas familial ties are patrilineal among peoples of the Eurasian steppes, as well as among the ancient Hungarians. This is why we focused on Y-chromosomal analyses to determine the genetic makeup of the people who arrived in the Carpathian Basin with the Hungarian tribes at the end of the ninth century CE. The ancient homeland Our analyses show that the paternal lineages of the Hungarian Conquerors are probably originated from three different larger regions far away from each other (see Fig. 11). Fig. 11 Altai-Lake Baikal-Tian Shan: “Altaic component” Five samples originate from this area, which is the farthest away from Hungary. The individual from Karos II grave 60 (C2-M86 haplogroup) is from the Turkic-speaking regions of Kazakhstan and the Altai Mountains. The individuals from Tuzsér grave 6 and Örménykút 52/50 (both N3a2-M2118) are from the region around Lake Baikal/Southern Siberia, where the Turkic-speaking Yakut also originate, whereas the individuals from Karos II grave 61 and Nagykőrös grave 2 (both R1a-Z93) are from South-Central Asia, near Turkic- and Iranian-speaking peoples. Although one of the R1a-Z93 samples is only a single step away (on 16 loci) from a Khazar sample in the median-joining network, the distribution of the Z93 branch (Rozhanskii and Klyosov 2012) and the nearby Kyrgyz hits point towards Central Asian ancestry. It is also possible that a Khazar descent joined the ancient Hungarian tribes in Khazaria. Olasz et al. (2018) studied the genetic origins (17-loci haplotype) of King Béla III of the Árpád dynasty and found that the paternal lineage of the Hungarian royal dynasty also belonged to Hg R1a. The authors did not conduct SNP testing; however, based on the haplotype data, the sample would likely belong to the Z93 branch. SNP testing should be done to verify our hypothesis. These five samples are still not enough to draw any far-reaching conclusions, but they do allow us to determine that this group of Hungarian Conquerors was likely represented by two or three tribes, which migrated separately west across Eurasia. According to Fettich (1935), the metalworks of the Hungarian Conquest period (and of the Avar period) were formed in the Minusinsk Depression in Southern Siberia. He believed that parallels of artifacts from the Hungarian Conquest period were found to the east and west of the Yenisei River, partly on the Abakan steppes, as well as in the areas surrounding Lake Baikal and the Altai Mountains, with the greatest amount in the Minusinsk Depression. His work remained almost completely without echo in the area of archaeology and history, which almost exclusively refers to the concept of the Uralian homeland. However, half a century later, anthropological studies also confirmed the importance of this geographical region in Hungarian prehistory. Tóth (1981) and Éry (1983) were the first in anthropology to recognize that some of the conquering Hungarians were of Asian origin. Anthropological studies based on craniometric comparisons of Hungarian Conquest period samples suggest that possible homeland of one part of the ancient Hungarian tribes, in addition to the abovementioned geographical regions, fall within South-Central Asia: Isfana, Fergana Valley, and the region of the Tian Shan (Fóthi 2014). Recent results in the field of archaeogenetics highlight the same regions as an important source of Hungarian ethnogenesis (Neparáczki et al. 2018, 2019). According to the median-joining network analysis of our study, one of the R1a-Z93 samples (Karos II grave 61) is from South-Central Asia (Tajik and Kyrgyz regions), and R1a-Z93 also dominates the paternal lineage of the Pashtuns living in the surrounding area (Underhill et al. 2015). The anthropological and subsequent genetic results will hopefully inspire further archaeogenetic research, beyond archaeology, history, and linguistics, to reveal the details of this still barely known area of Hungarian prehistory. The results of our genetic study are in agreement with the anthropological results outlined above. Both the calculated STR-based age of the N3a2 subgroup (1970–2070 ± 690 years; i.e., the period of the Asian Huns) and the geographic location of the homologous cases suggest that these samples may have belonged to the Hun/Turkish line of the Hungarian Conquerors. The two N3a2 and C2-M68 Hungarian Conqueror warriors may have originated from the area between Lake Baikal and the Altai Mountains, while the R1a-Z93 warriors may have been from the Tian Shan in South-Central Asia.The widespread area from the Altai Mountains to the Tian Shan reaches the edge of the Asiatic Hunnic/Xiongnu territory. Several skeletal remains of Asiatic Huns/Xiongnu have been discovered, such as from the Egyin Gol cemetery with haplogroups Q-M242, N1c, (N3a in modern terminology), and C3-M130 (Keyser-Tracqui et al. 2003; Kang et al. 2013) and the Duurlig Nars site with haplogroups: C3-M130 and R1a1 (Kim et al. 2010) in Northern Mongolia. To the southern part of the Asiatic Huns’ territory (China), only males with Q haplogroups were found in the Xiongnu cemeteries. In Xinjiang (Barköl Kazakh Autonomous County), the 13 examined ancient individuals belonged to haplogroup Q (Q-M346, Q*, Q1a, and Q1b) (Kang et al. 2013), and in Pengyang, each of the 4 examined skeletons also belonged to haplogroup Q (Zhao et al. 2010). We found 2 haplogroups among the Hungarian Conqueror skeletal remains, which are present in Southern Siberian and Central Asian samples of the Xiongnu period (R1a and N3a2), but there were no Q haplogroup individuals in the research sample. Damgaard et al. (2018) describe a Y-DNA and autosomal genetic shift in the Baikal region between the Baikal Early Neolithic (Y-Hg N and a mix of Ancient North Eurasian and Ancient East Asian autosomal components) and the Baikal Late Neolithic/Early Bronze Age (arrival of Y-Hg Q and additional Ancient East Asian components). This supports the assumption that proto-Uralic populations migrated to the West and were replaced by possible proto-Altaic populations during the 1500-year period that separates the Baikal Neolithic from the Early Bronze Age which would imply that later Xiongnu/Huns spoke a Turkic language through autosomal aDNA analysis.
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