NEW paper on old Magyars!

**Jana** · 07-15-2020, 11:07 AM

https://www.biorxiv.org/content/10.1...07.13.200154v1

The ancient Hungarians originated from the Ural region of Russia, and migrated through the Middle-Volga region and the Eastern European steppe into the Carpathian Basin during the 9th century AD. Their Homeland was probably in the southern Trans-Ural region, where the Kushnarenkovo culture disseminated. In the Cis-Ural region Lomovatovo and Nevolino cultures are archaeologically related to ancient Hungarians. In this study we describe maternal and paternal lineages of 36 individuals from these regions and nine Hungarian Conquest period individuals from today's Hungary, as well as shallow shotgun genome data from the Trans-Uralic Uyelgi cemetery. We point out the genetic continuity between the three chronological horizons of Uyelgi cemetery, which was a burial place of a rather endogamous population. Using phylogenetic and population genetic analyses we demonstrate the genetic connection between Trans-, Cis-Ural and the Carpathian Basin on various levels. The analyses of this new Uralic dataset fill a gap of population genetic research of Eurasia, and reshape the conclusions previously drawn from 10-11th century ancient mitogenomes and Y-chromosomes from Hungary.

More support for Ugric/Uralic oldest base of Hungarians.

**Jana** · 07-15-2020, 11:13 AM

The Ural region was involved in numerous migrations, which events also shaped the history of
53 Europe. The archaeological imprint of these events can be witnessed among others on the early
54 medieval cemeteries of the South-Ural region. Compact cemeteries with few hundred tombs
55 are typical of this territory, which have provided rich archaeological findings first in the last
10-15 years1–5
56 . According to archaeological, linguistic and historical arguments, the
ethnogenesis of modern Hungarian population can be traced back to the Ural region1,6,7 57 .
58
59 Based on linguistic evidences, the Hungarian language, belonging to the Ugric branch of the
60 Uralic language family, was developed at the eastern side of Ural Mountains between 1000-
500 BC8,9 61 . According to the written and linguistic sources and archaeological arguments, after
the 6th 62 century AD, part of the predecessors of Hungarians moved to the Western Urals (CisUral region) from their ancient homeland. Around the first third of 9th 63 century AD a part of this
64 Cis-Uralic population crossed the Volga-river and settled near to the Khazarian Khaganate in
the Dnieper-Dniester region1–5,10 65 (Fig. 1). Early Hungarians lived in Eastern Europe (forming
66 the so-called Subbotsy archaeological horizon) until the conquest of the Carpathian Basin that
took place in 895 AD. The material traits of 10th 67 century AD Carpathian Basin was rapidly
68 transformed after the conquest, its maintained cultural connections with East-European regions
have numerous doubtless archaeological evidence2,4,11 69 .
70
71 Genetic history of prehistoric to medieval populations of the Ural region have been scarcely
72 investigated to date. On the other side, the populations of the medieval Carpathian Basin have
been intensively studied from the perspective of uniparental markers12,13 73 . Recently, Neparáczki
74 et al. have published 102 whole mitogenomes from early Conquest period cemeteries in
Hungary14 75 . Authors have suggested that the mixed population of steppe nomads (Central Asian
76 Scythians) and descendants of the East European Srubnaya culture’s population among other
77 undescribed populations could have been the basis of genetic makeup of Hungarian conquerors.
Their results furthermore assume Asian Hunnic-Hungarian conqueror genetic connections14 78 . It
79 is important to note, that the investigated medieval sample set does not represent the conqueror
80 population as a whole, hence 76% of the samples originated from a special site complex Karos81 Eperjesszög from northeast Hungary, which is one of the most important sites of the Hungarian
82 Conquest period with many findings of eastern characteristics as well. The conclusions are
83 large-scale, but the most highlighted connection with the population of the Srubnaya culture is
84 vague, because it existed more than 2000 years before the appearance of the first traces of
85 ancient Hungarians’ archaeological heritage. Additionally, further mentioned relations such as
86 the Xiongnu (Hunnic) genetic dataset is bare from Eurasia, and Huns’ genetic heritage is
87 basically unknown, as well.
88 Two recent articles have investigated the Y-haplogroup variability of Hungarian conquerors
89 describing the conqueror´s elite population as heterogenous, with significant proportion of
European, Finno-Permic, Caucasian and Siberian (or East Eurasian) paternal lineages15,16 90 . Fóthi
91 et al. have claimed that the Hungarian conquerors originated from three distant sources: Inner
92 Asia (Lake Baikal – Altai Mountains), Western Siberia – Southern Urals (Finno-Ugric peoples)
93 and the Black Sea – Northern Caucasus (Northern Caucasian Turks, Alans, and Eastern
Europeans)15. Both studies15,16 94 pointed out the presence of the Y-haplogroup N-Z1936 (also
95 known as N3a4-Z1936 under N-Tat/M46), which is frequent among Finno-Ugric speaking
peoples17 96 . This lineage also occurs among modern Hungarians in a frequency up to 4%. Post et
97 al. have reconstructed the detailed phylogeny of N-Z1936 Y-haplogroup showing that specific
98 sublineages are shared by certain ethnic groups, e.g. N-Y24365/B545 by Tatars, Bashkirs and
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3
99 Hungarians, which connect modern-day Hungarians to the people living in the Volga-Ural
region17 100 .
101 Earlier mitochondrial DNA (mtDNA) studies of modern populations speaking Uralic languages
102 suggest that the distribution of Eastern and Western Eurasian mtDNA lineages are determined
by geographic distances rather than linguistic barriers18–20 103 , e.g. Finno-Ugric populations from
104 Volga-Ural region seem to be more similar to their Turkic neighbours than to linguistically
related Balto-Finnish ethnic groups18 105 . The recent study of 15 Uralic-speaking populations
106 describes their similarities to neighbouring populations as well, however they also share genetic
component of possibly Siberian origin21 107 . In spite of the unambiguously Central-European
characteristics in mtDNA makeup12,22 108 , this statement also can be applied to modern day
Hungarians23 109 .
110
111 The main goal of this study is to expand the current set of archaeological knowledge about the
112 early medieval populations of the Ural region by archaeogenetic methods. During the collection
113 of 36 samples from Ural region processed in this study, the most important intention was to
114 collect samples exclusively from such professionally excavated and appropriately documented
115 cemeteries from the South-Ural region, which are culturally and temporally (directly or
116 indirectly) connected to the ancestors of Hungarians (Fig.1 and Supplementary Figs. S1a-h).
117 The sampled Uyelgi cemetery from Trans-Ural region presented the greatest similarity to the
118 archaeological traits of the tenth-century Carpathian Basin (Figs. 1-2, Supplementary Figs. S1eh). This cemetery of the late Kushnarenkovo culture was used between the end of 8th 119 century to
11th century2,24 120 .
121 As the archaeological and historical theories are slightly diverse, we aimed to cover a wide
122 range of early medieval archaeological cultures located in the middle course of the Kama river
123 in the west side of the Ural Mountains (Cis-Ural region). Scholars connect the termination of
the Nevolino Culture in 8-9
th 124 centuries AD to the westward migration of ancestors of
Hungarians1–3
125 , hence the sampling was carried out in all three phases of this culture: Brody
(3rd
-4
th centuries), Bartym (5-6
th centuries) and Sukhoy Log (7-8
th centuries)25 126 (Fig. 1).
Furthermore, we investigated the Bayanovo cemetery (9-10th 127 centuries AD), which represents
the southern variant of Lomovatovo culture3
128 that shows close cultural connection to its southern
129 neighbour Nevolino culture. The sampling of the richly furnished graves of Bayanovo was
limited by the poor preservation of bone samples (see Supplementary text, Figs S1b-d)6
130 .
131 Additionally, we reanalysed nine samples from tenth- to twelfth-centuries ancient Hungarians
132 for whole mitogenomes from the Carpathian Basin, who were chosen from the previous study
Csősz et al.13 133 based on identical hypervariable I region (HVRI) haplotypes of mtDNA with
134 some of investigated Uralic individuals.
135 In this paper, our main purpose was to characterize the maternal and paternal genetic
136 composition of populations from the third- to eleventh-centuries South-Ural region and
137 compare the results with the available ancient and modern genetic datasets of Eurasia. We also
138 aimed to describe possible genetic connections between the studied Uralic populations and the
139 Conquest period populations of the Carpathian Basin.

**Jana** · 07-15-2020, 11:15 AM

Conclusion

The Ural region had an important role in ancient Hungarians’ ethnogenesis based on
389 archaeological, linguistic and historical sources, although the results of these research fields
390 exhibit differences of chronological and cultural aspects. The here presented new mitogenome,
391 Y-chromosome and shallow shotgun autosomal DNA sequence data from the South-Urals
392 confirms the region’s relevance from population genetic perspective too.
393 The overall maternal makeup of the investigated 36 samples from the Ural region in a
394 phylogenetic and phylogeographic point of view suggests a mixed characteristic of rather
395 western and rather eastern components, although the paternal lineages are more homogenous
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396 with Y-haplogroups typical for the Volga-Ural region. The exact assignment of each
397 mitochondrial haplotype of the Trans-Uralic Uyelgi population to the eastern and western
398 Eurasian components is impossible, but comprehensive representatives are present.
399 Mitochondrial haplogroups of European origin N1a1a1a1a and H40b provide a horizon-through
400 success of maternal lineages with inner diversification, which suggests a base population of a
401 rather western characteristics. On the other hand, identical (C4a1a6) or single (A, A12a,
402 C4a2a1) haplotypes with strong eastern phylogeography, highly pronounced in the third
403 horizon, suggest a relatively recent admixture to this population. The apparent co-occurrence
404 of genetic and archaeological shift is however contradicted by the homogeneity of ancestry
405 components, nuclear genomic PCA positions, homogeneity of paternal makeup (although this
406 one itself can be explained by patrilocality), and presence of eastern component (C4a1a6) in all
407 horizons. Despite the fact that the genetic contribution of a population related to the Srostki
408 culture cannot be excluded at this level, it is more likely that the majority of eastern components
409 admixed before the usage of the Uyelgi cemetery. The uniparental genetic composition of
410 Uyelgi population signals them as a chronologically and/or geographically related population
411 to the possible genetic source of the Hungarian conquerors. Furthermore, their preliminary
412 autosomal results show that they shared their allele frequency makeup with modern Uralic and
413 West Siberian populations that are linguistically or historically related to Hungarians, which
414 provide a good standpoint for future studies.
415 The maternal phylogenetic connections of Uyelgi with Hungarian conquerors can be divided to
416 indirect (monophyletic but not successive) and direct (identical or one-step neighbour)
417 relationships. Interestingly, indirect connections can be genetically assigned to the western418 characteristic base population, whereas direct connections are almost exclusive to the admixed
419 eastern component. One possible explanation for this phenomenon is that Hungarian
420 conquerors and Uyelgi shared common ancestry in the past that separated prior eastern
421 admixture, latter which provided genetic components subsequently to both groups. The exact
422 origin or identification of the eastern component yet to be described, however, nuclear
423 admixture proportions and loose phylogenetic connections points towards Central Asia, but
424 further and deeper analyses with extended dataset is required for firm our statements.
425 The phylogenetic makeup of Cis-Ural region questions their compactness or successiveness;
426 however, the scarce data does not allow extensive analysis for this group. Hungarian conqueror
427 connections here are sporadic, but regional affinity is observable, which is more pronounced in
428 MDS and PCA. Earlier studies based solely on the genetic makeup of Hungarian conquerors
429 tend to connect the non-European lineages to various eastern regions, but especially the
430 presence of rare Far East haplotypes in the Late Iron Age and Early Medieval Cis-Ural group
431 may reshape these conclusions in the future.

**Jana** · 07-15-2020, 11:17 AM

Primary observations

153
154 45 high coverage mitochondrial genomes were obtained (sequencing depth from 8.71× to
155 154.03×), with mean coverage of 71.16× and an average contamination rate of 0,2%. The new
156 dataset consists of the mixture of nine macrohaplogroups (A, C, D, H, T, U, N, R, Z) (Fig. 3a).
157 Haplogroups of presumably west Eurasian origin are represented by U (U2e1, U3a1, U4a1d,
158 U4b1a1a1, U4d2, U5a1a1, U5b2a1a1, N=12), H (H1b2, H3b, H40b, N=9), N (N1a1a1a1a,
159 N=5) and T (T1a1, T1a2, T2b4h, N=5), although phylogeographic analyses show eastern origin
160 for some of them, see Table1 and Supplementary Figs. S4a-s. Eastern Eurasian lineages are
161 represented by A (A+152+16362, A12a, N=4), C (C4a1a6, C4a2a1, N=6), D (D4j, D4j2, N=2),
162 along with R11b1b and Z1a1a by one individual each (Fig. 3a).
163
164 Even though that the Hungarian conquerors were selected based on mtDNA HVRI matches
165 with certain ancient individuals from the Ural region, they have not proved to be identical on
166 whole mitogenome level, but remained phylogenetically close to the associated samples (see
167 Supplementary Figs. S4a-s).
168 A few mitochondrial lineage relations connect Trans-Ural and Cis-Ural regions: e.g. samples
169 from Uyelgi and Sukhoy Log clustered together in one main branch of the A+152+16362
170 haplogroup tree (Supplementary Fig. S4b), furthermore samples from Uyelgi and Bartym (with
171 haplogroup U4d2) are located on the same main branch as well (Supplementary Fig. S4p).
172 The sole investigated sample from Brody cemetery with haplogroup D4j2 neither show close
173 maternal genetic connection to other Uralic samples nor to Hungarian conquerors.
174 In contrast to the mitochondrial lineages, the Y-chromosomal gene pool based on STR and/or
175 SNP data show homogenous composition in our dataset: 83.3% is N-M46, 5.5% G2a (G176 L1266), 5.5% J2 and 5.5% is R1b of the typed male individuals (Supplementary Table S2). 13
177 male samples out of 19 from Uyelgi cemetery carry Y-haplogroup N with various DNA
178 preservation-dependent subhaplogroup classifications, while in the Cis-Ural we detected three
179 N-M46 Y-haplogroups (samples from Brody, Bartym and Bayanovo cemeteries). The overall
180 poor preservation of further Cis-Uralic samples from Sukhoy Log and Bartym disabled further
181 Y-chromosome-based analyses (Supplementary Table S2).
182

183 Comparative population genetic analyses of maternal lineages and genomic data

184
185 We performed population genetic statistical analyses as well. The principal component analysis
186 (PCA) and Ward clustering of 50 ancient and 64 modern populations were performed separately
187 (Fig. 3b, Supplementary Figs. S5-S8), based on haplogroup frequencies (Supplementary Tables
188 S3 and S4). The Hungarian conquerors are the closest population to the Cis-Ural group on the
189 PCA (along PC1 and PC2 components, see Fig. 3b) and this population is relatively near to the
190 Uyelgi among the Iron Age population from Central-Asia and the East European Scythians
191 along PC1 and PC3 components (Supplementary Fig. S5), because these ancient populations
192 have mixed pool of western and eastern Eurasian macrohaplogroups, which is unusual in
193 European and Asian populations that are separated along the PC1. The nearby position of Cis194 Ural and Uyelgi to the Hungarian conquerors is displayed on the mtDNA haplogroup-based
195 Ward type clustering tree too, where they appear in the same main branch (Supplementary Fig.
196 S6). Some of Central-South Asian and Finno-Ugric modern populations (e.g. Khanty and
197 Mansi) show close connections to the investigated Cis-Ural and Uyelgi populations based on
198 Ward-type clustering and PCA (Supplementary Figs. S7 and S8). The haplogroup frequencies
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199 of three highlighted populations are displayed on the Fig.3a diagram. The mitochondrial
200 haplogroup pool of the Hungarian conquerors´ large sample-set is the most diversified and
201 contains nearly all haplogroups obtained in two populations from Ural region with a similar
202 proportion of haplogroups with western and eastern Eurasian origin. This phenomenon causes
203 their relatively nearby positions on the PCA and Ward clustering tree.
204 Pairwise FST values of populations indicate non-significant differences of the Cis-Ural from 13
ancient populations (Supplementary Table S5), among them the Hungarian conquerors14 205 show
206 the lowest genetic distance (FST = 0.00224) (for further FST values, p values, and references see
207 Supplementary Table S5). According to the MDS plot of 28 ancient populations based on
208 linearized Slatkin FST (Supplementary Fig. S9a), the Cis-Ural population shows affinities
209 among others to the populations of medieval Hungarian conquerors along coordinates 1 and 2,
210 and is situated between European and Asian populations, which reflects the raw FST values. The
211 Uyelgi is standing on the Asian part of the plot relatively far from all ancient populations, which
212 is most likely due to its significant and larger genetic distances from ancient populations (except
the Late Iron Age population from Central Asia26 213 ) and the scarcity of Asian comparative
214 mitogenome datasets. The rank correlation heatmap (Supplementary Fig. S9b) of the FST values
215 of ancient populations supports the MDS plot, where the Uyelgi and Cis-Ural populations
216 cluster with the same ancient populations that are close to them on the MDS plot.
The genetic connection of Cis-Ural population and Hungarian conquerors14 217 is obvious based
218 on pairwise FST calculation and is visible on the PCA and MDS plots as well, where they are
219 the closest, although direct phylogenetic connections are scarce. This indicates geographical
proximity of their former settlement area, rather than a direct connection. Neparáczki et al.14 220
221 have described the Hungarian conqueror mitogenome diversity in essence as a mixture of
222 Srubnaya and Asian nomadic populations. Their analyses and interpretation were restricted by
223 the lack of ancient samples from the Ural region, whereas new data now refine such previous
conclusions14 224 . Furthermore, it is notable, that the previously studied Hungarian conqueror
225 population is a pool of mixed origin including not only immigrants but also local admixed
226 lineages from the Carpathian Basin.
227 The Cis-Ural population reveals non-significant genetic distances from four modern
228 populations of Central Asian Highlands, furthermore seven populations of Near East and
229 Caucasus region and six European populations (see Supplementary Table S6) indicating a
230 mixed character of this population, which is also visible on the MDS plot.
231 Interestingly, the mitogenome pool of Uyelgi shows significant differences in genetic distances
232 among nearly all prehistoric and modern populations including Hungarian conqueror
233 population in spite of the extensive phylogenetic connections, which might be explained by
234 high amount of related lineages within the population, as well as by their mixed character of
235 Eastern- and Western-Eurasian haplogroups.
236 We performed genomic PCA of five Uyelgi samples consisting of 10,828 nuclear genomic
237 SNPs on average gained from 3000 SNP capture and shallow shotgun sequencing data (from
238 598,094 called SNPs). The five samples are plotted together on the genomic PCA and they also
239 appear close to the modern Bashkir and Siberian Tatar individuals as well as to the Altaian
Bronze Age Okunevo population27, to a hunter-gatherer individual from Tyumen region28 240 and
Iron Age Central Sakas from Kazakhstan26 241 (see Supplementary Information, chapter 3 and
242 Supplementary Figs. S3a-c) in line with the uniparental makeup. Since PCA may not reveal
243 population stratification we performed unsupervised ADMIXTURE (K=16) on an enlarged sets
244 of SNPs (SI, chapter 3). The five Uyelgi samples with an average calling of 22,540 SNPs show
245 the most similar ancestry cluster proportions to present-day Mansis and Irtysh-Barabinsk Tatars
and to a set of various populations lived in the Central Steppe region27 246 .
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247 To disentangle the connections between these populations and possible population genetic
248 events of thousands of years between populations under study, more ancient reference samples
249 and deeper sequencing for more detailed analyses are needed.

250
251 The genetic continuity between the horizons of the Uyelgi cemetery (Trans-Ural region)

252
253 The kurgan burials at Uyelgi site can be divided into at least three chronological horizons:
254 I.) the oldest ninth-century, II.) ninth- and tenth-centuries and III.) tenth- and eleventh-centuries
255 according to the archaeological records (see Supplementary text chapter 1 and Supplementary
256 Figs. S1e-h). Uniparental genetic markers show genetic continuity between these horizons
257 suggesting maternally rather endogamous population, which could not be observed in
258 archaeological findings due to high number of disturbed burials in the cemetery. Mitochondrial
259 phylogenies of N1a1a1a1a, C4a1a6 and H40b provide identical or monophyletic lineages
260 within and between the three horizons (see Figs. 4-5 and Supplementary Figs. S4h-g), which
261 trend is more pronounced by haplotype and network analysis of paternal lineages (Fig. 6.,
262 Supplementary Figs S11-12).
263 The haplotypes of N-M46 Y-haplogroup are presented in all three horizons, however with little
264 differences in STR profiles (Supplementary Table S2). The oldest and the middle horizons
contain only N-M46 haplotypes including two identical STR profiles in Kurgan 32 (9th 265 century).
266 Three identical Y-STR profiles are detected among individuals of Kurgans 28, 29 and 30 (Fig.
267 4 and Fig. 6). Probably further identical Y-haplotypes could have been in this cemetery, but the
268 preservation has not let us reconstruct whole Y-STR profiles of seven males (see
269 Supplementary Table S2). Based on these results we suggest that Uyelgi cemetery was used by
270 a patrilocal community.
The genetic continuity between the 9–11th 271 centuries is also supported by genomic data
(Supplementary Figs. S3a-c). The Uyelgi2 sample of the youngest horizon (10–11th 272 centuries)
has high proportion of shared drift with the Uyelgi10 of the 9–10th 273 centuries.
274

275 The possible maternal genetic connection of South-Ural region’s populations and the
276 Hungarian conquerors

277
The genetic connection of Uyelgi cemetery in the Trans-Ural and 10th 278 century Hungarian
279 conquerors in the Carpathian Basin is supposed by close maternal relationships of the following
280 individuals: Uyelgi3 from Kurgan 28 of the youngest horizon and three Hungarian conquerors
from Karos II cemetery14 281 have identical U4d2 mitogenome haplotype (Supplementary Fig.
282 S4p). Furthermore, the mtDNA A12a lineage of Hconq3 (30-40 years old woman from Harta
cemetery dated to the first half of 10th 283 century AD) is an ancestor of the mtDNA lineage of
284 Uyelgi7 (from Kurgan 30 of the youngest horizon of the cemetery) based on the A12a
285 haplogroup tree (see Supplementary Fig. S4a).
286 The mentioned graves from Uylegi show the characteristic of the Srostki culture, where the gilt
287 silver mounts with plant ornaments were typical, and which was disseminated from the Siberian
288 Minusinsk Depression and the Altai region through the Baraba Steppe and North-Kazakhstan
289 to the Trans-Ural region (Fig. 1). Moreover, it is notable that the archaeological findings in
these kurgans are dated not earlier then the10th 290 century AD, i.e. after the Hungarian conquest
291 of the Carpathian Basin. The Hungarian conquerors from Karos cemetery appearing on these
292 phylogenetic trees could represent the first generation of conquering populations based on their
293 grave material, therefore identical mitogenome sequences can point out close biological
294 connections or common source population of the Uyelgi population and the Hungarian
295 conquerors.
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296 The D4j phylogenetic tree contains one interesting phenomenon: the mitochondrial lineage of
297 the sample Uyelgi21 from the Kurgan 11 located in the oldest horizon of Uyelgi cemetery
298 clusters only with one modern-day Hungarians, whose lineage is ancestral to the lineage of
299 Uylegi21. The findings of this Kurgan 11 (belonging to the Srostki culture) show similarities
300 to the typical findings of the Hungarian conquerors from the Carpathian Basin as well (see
301 Fig. 2 and Supplementary Fig. S1h).
302 The mitogenome of individual Uyelgi10 and three identical lineages of two Hungarian
303 conquerors (Hconq1 and Hconq6) from Balatonújlak-Erdő-dűlő and Hconq9 from Makó-Igási
304 járandó cemetery clustered together in one branch on the phylogenetic tree of haplogroup
305 U5a1a1 (Supplementary Fig. S4q). The Uyelgi10 from Kurgan 7 of the middle horizon of the
306 cemetery shows mixed character from archaeological point of view: the findings can be
connected to the 9th 307 century AD as well as to the cultural influences of the Srostki culture (for
the detailed information see Supplementary information)29,30 308 . The samples of adult women
309 from Balatonújlak-Erdő-dűlő buried with gilt silver hairpins could be dated (based on
archaeological findings) to the middle third of the 10th century AD31 310 . One of their burials had a
311 grave with a sidewall niche of eastern origin. The grave from Makó-Igási járandó without
findings is dated to the middle third of 11th 312 century AD, i.e. to the Árpádian Age, when
313 conquerors and the local population presumably admixed already. Interestingly, the 25-30 years
old man shows some Asian cranial traits as the most men buried in this cemetery32 314 .
315 The connection of Uyelgi cemetery and Hungarian conquerors is visible on the N1a1a1a1a
316 branch of the tree of haplogroup N1a1 too, that was prevalent among the ancient Hungarians
317 (Fig. 5). Here seven Hungarian conqueror samples from cemeteries Kenézlő-Fazekaszug,
318 Orosháza-Görbicstanya and Karos-Eperjesszög clustered together on one branch, while the five
319 Uyelgi samples from the earliest and latest horizons are located together next to this branch.
320 These results signalize indirect connection between these two populations and don’t speak for
321 their direct successiveness but rather for their common source in agreement with the
322 archaeological chronology of Uylegi site.
323 The maternal genetic connection of the Cis-Ural region and the Hungarian conquerors is
324 apparent especially on the phylogenetic tree of mitochondrial haplogroup T2b4h, where
Bartym2, Bay3 and Hungarian conqueror from Karos site14 325 are located on the same branch,
326 moreover, the individuals from Bartym and Karos share the same lineage that is ancestral to the
327 mtDNA lineages of individual from Bayanovo (Supplementary Fig. S4k). The lineage of Karos
(K1/3286) sample was determined as of possibly Asian origin by Neparáczki et al.14 328 ,
329 nevertheless, their assumption is revisited by our data, not only by actual phylogenetic
330 connections but due to the recurrent western presence of eastern lineages even from pre331 medieval times. The burial of this adult male in Karos was without findings because disturbance
332 of the Karos I cemetery’s burials by agricultural activity.
333

334 Ancient paternal lineages of the South-Ural region

335
336 Majority of Uyelgi males belonged to Y chromosome haplogroup N, and according to combined
337 STR, SNP and Network analyses they belong to the same subclade within N-M46 (also known
338 as N-tat and N1a1-M46 in ISOGG 14.255). N-M46 nowadays is a geographically widely
distributed paternal lineage from East of Siberia to Scandinavia33 339 . One of its subclades is
340 N-Z1936 (also known as N3a4 and N1a1a1a1a2 in ISOGG 14.255), which is prominent among
341 Uralic speaking populations, probably originated from the Ural region as well and mainly
342 distributed from the West of Ural Mountains to Scandinavia (Finland). Seven samples of Uyelgi
343 site most probably belong to N-Y24365 (also known as N-B545 and N1a1a1a1a2a1c2 in
344 ISOGG 14.255) under N-Z1936, a specific subclade that can be found almost exclusively in
todays’ Tatarstan, Bashkortostan and Hungary17 345 (ISOGG, Yfull).
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346 Median Joining (MJ) network analysis is performed using 238 N-M46 Y-haplotypes including
347 seven samples from Uyelgi detected with 17 STR loci (Fig. 6, Supplementary Table S8) as well
348 as 335 N-M46 Y-haplotypes with 12 STR loci (Supplementary Fig. S12, Supplementary Table
349 S8). Based on MJ of 17 Y-STR loci, certain samples show identical or one-step neighbour
profiles to Bashkirs, Khantys17, Hungarians34 350 , Tatars from Volga-Ural region and a Central
Russian sample17 351 (Fig. 6). The MJ based on 12 Y-STR data show one-step neighbour
352 connection of Uylegi with two Hungarian conquerors from Bodrogszerdahely-Bálványhegy
and Karos-Eperjesszög15 353 (Supplementary Fig. S12). YHRD online database show further
354 affinities or identities among Finnish, Ural region (Sverdlovsk Oblast) or European Russian
355 region (Penza and Arkhangelsk Oblasts) samples, notably either from territories of Uralic
356 language affinities or along the supposed migration route of early Hungarians. It is noteworthy
that the seventh-century Avar elite from the Carpathian Basin35 357 , in spite of the similar N-M46
358 frequency to Uyelgi, had a distant subtype (N-F4205, N1a1a1a1a3a in ISOGG 14.255), which
is prominent in present-day Mongolic speaking populations around Lake Baikal33 359 . Furthermore
360 they had a fairly different population history than populations of this study, therefore they shall
not be confused with each other35 361 .
362
363 Uyelgi11 from Kurgan 29 belongs to J2 Y-haplogroup. The Y-haplogroup J is widespread
nowadays descended from the Near East36 364 . Interestingly, a Hungarian conqueror from
Sárrétudvari-Hízóföld (SH/81) carries the J2a1a subgroup16 365 , however Uyelgi11 could not be
366 typed downstream to J2 and therefore further assumptions cannot be made at this level.
367
368 Uyelgi4 belongs to G-L1266 (G2a2b2a1a1a1b in ISOGG 14.255), which sublineage is
369 confirmed to be present outside of Europe within the European G-L140 branch of G. Among
370 Hungarian conquerors the presence of G-L30 (G2a2b in ISOGG 14.255) was attested by
Neparáczki et al.16 371 from Karos II (K2/33) without further classification or STR data, but
recently G-L1266 is confirmed by Fóthi et al.15 372 which sample could also be included in our
373 STR analysis. By using 14 STR markers in this case, due to the limitations of the database,
374 MJ network shows a Caucasian affinity of both Hungarian conqueror (RP/2) and Uyelgi
375 individuals (Supplementary Fig. S11, Supplementary Table S9), however, neither identity nor
376 monophyly can be observed between them.
Both studies15,16 377 indicate Caucasian origin for part of the Hungarian conquerors based on the
378 prevalence of this specific G2a Y-haplogroup. This hypothesis cannot be confidently excluded
379 by our data nor our network analysis, however its presence in Uyelgi site could reshape this
380 theory in the future.
381 In the Cis-Ural sample set the DNA preservation was insufficient for proper paternal lineage
382 analyses, the only obtained N-M46 Y-haplotype of Bay2 sample and the R1b haplotype of
383 Bartym3 do not have direct matches in the worldwide YHRD database, however, we found four
384 one-step-neighbours of Bay2 from Sverdlovsk Oblast (Ural region) and Lithuania.

**vbnetkhio** · 07-15-2020, 11:38 AM

Originally Posted by Feiichy

https://www.biorxiv.org/content/10.1...07.13.200154v1

More support for Ugric/Uralic oldest base of Hungarians.

i think there have been 4 or 5 of Magyar Y-DNA papers already in the past few years? it would be interesting to put all those haplogroups together and check where their closest modern matches on yfull cluster.

**Jana** · 07-15-2020, 11:54 AM

Originally Posted by vbnetkhio

i think there have been 4 or 5 of Magyar Y-DNA papers already in the past few years? it would be interesting to put all those haplogroups together and check where their closest modern matches on yfull cluster.

I agree! It's interesting how this paper implies recently published studies on conquerors had some flaws:

Authors have suggested that the mixed population of steppe nomads (Central Asian Scythians) and descendants of the East European Srubnaya culture’s population among other undescribed populations could have been the basis of genetic makeup of Hungarian conquerors. Their results furthermore assume Asian Hunnic-Hungarian conqueror genetic connections . It is important to note, that the investigated medieval sample set does not represent the conqueror population as a whole, hence 76% of the samples originated from a special site complex Karos Eperjesszög from northeast Hungary, which is one of the most important sites of the Hungarian Conquest period with many findings of eastern characteristics as well. The conclusions are large-scale, but the most highlighted connection with the population of the Srubnaya culture is vague, because it existed more than 2000 years before the appearance of the first traces of ancient Hungarians’ archaeological heritage. Additionally, further mentioned relations such as the Xiongnu (Hunnic) genetic dataset is bare from Eurasia, and Huns’ genetic heritage is basically unknown, as well.

Neparáczki et al.14 2020 have described the Hungarian conqueror mitogenome diversity in essence as a mixture of Srubnaya and Asian nomadic populations. Their analyses and interpretation were restricted by the lack of ancient samples from the Ural region, whereas new data now refine such previous conclusions . Furthermore, it is notable, that the previously studied Hungarian conqueror population is a pool of mixed origin including not only immigrants but also local admixed lineages from the Carpathian Basin.