Alans and Slavs

[Vojvoda]

Alan tribes living north of the Black Sea may have moved northwest into what is now Poland, merging with Slavic peoples there to become the precursors of historic Slav nations (notably Serbs and Croats). Third-century inscriptions from Tanais, a town on the Don River in modern Ukraine, mention a nearby Alan tribe called the Choroatos or Chorouatos. The historian Ptolemy identifies the 'Serboi' as a Sarmatian tribe who lived north of the Caucasus, and other sources identify the Serboi as an Alan tribe in the Volga-Don steppe in the third century.

Accounts of these names reappear in the fifth century, with the Serboi, or Serbs, established east of the river Elbe in what is now western Poland, and the Croats in what is now Polish Galicia. The Alan tribes likely moved northeast and settled among the Slavs, dominating and mobilizing the Slavic tribes they encountered and later assimilating into the Slav population. In 620 the Croats and Serbs were invited into the Balkans by Eastern Roman Emperor Heraclius to drive away the Turkic Avars, and settled there among earlier Slavic migrants to become ancestors of the modern Serbs and Croats. Some Serbs remained on the Elbe, and their descendants are the modern Sorbs. Tenth-century Byzantine and Arab accounts describe a people called the Belochrobati (White Croats) living on the upper Vistula, an area later called Chrobatia.

[Menydh]

Weren't Alans an Indo-European tribe? (more "indos-", mind you).

[Carnyx]

I read they might have been associated with indo-iranians.

[Zrinski]

Alans are sub-ethnos of old Iranic tribes. Namely the Sarmatians and Schytians.

Hello our Ossetian brothers.

[bocian]

There are definite links between Alans/Sarmatians/Scythians and modern day Slavic populations. It's not like they disappeared into thin air.

There are genetic and traditional similarities between Poles and Sarmatians/Alans.

Sarmatian Tamgas and Polish coats of arms are strikingly similar, the long tradition of Polish horsemanship (quite odd if Slavic ethnogenesis is in supposed marshes ) and this study:


(I've posted this before, thanks norda)




Genetic Analysis of a Scytho-Siberian Skeleton and Its Implications for Ancient Central Asian Migrations




Abstract The excavation of a frozen grave on the Kizil site (dated to be 2500 years old) in the Altai Republic (Central Asia) revealed a skeleton belonging to the Scytho-Siberian population. DNA was extracted from a bone sample and analyzed by autosomal STRs (short tandem repeats) and by sequencing the hypervariable region I (HV1) of the mitochondrial DNA. The resulting STR profile, mitochondrial haplotype, and haplogroup were compared with data from modern Eurasian and northern native American populations and were found only in European populations historically influenced by ancient nomadic tribes of Central Asia.

KEY WORDS: ANCIENT DNA, HV1 SEQUENCE, STRS, HAPLOGROUPS, SCYTHIAN POPULATION, CENTRAL ASIA, ALTAI REPUBLIC, KIZIL SKELETON (95-KBI-52), MIGRATION, MOLECULAR ARCHEOLOGY, D3S1358, VWA, FGA, D8S1179, D21S11, D18S51, D5S818, D13S317, D7S820, AMELOGENIN

The Central Asian region has been inhabited since the lower Paleolithic Era and is considered a crossroads of migration routes of ancient nomadic populations. During the Neolithic period, Central Asia was a contact area between nomadic tribes and the neighboring agriculturist populations. The Scythians (700 B.C.-A.D. 200) were one of the most famous nomadic populations living in the Eurasian steppe zone. The Scytho-Siberian group, localized in southern Siberia, composed the eastern extremity of this population.



The history of the Scythians is known only from ancient texts (Achaemenid, Greek, and Chinese sources) and by the excavation of their burial places. The archeological records (Bokovenko 1994; Dvornichenko and Fedorov-Davydov 1994; Francfort et al. 2000) and morphological (Chikisheva 2000a,b) and genetic (Clisson et al. 2002; Voevoda et al. 2000) studies of this ancient population have not led to a conclusive scenario regarding the origin and fate of the Scythians. Only an increased number of ancient nomad grave discoveries, their further study, and biological analysis of the human remains will allow for a better understanding of the origins, migrations, and disappearance of the Scythian populations.

In this study, two types of genetic markers (HVl region of the mtDNA and autosomal STRs) were used to analyze DNA recovered from the remains of a 2500-year-old human skeleton excavated from the Kizil valley (Altai mountains). Autosomal STRs are used in ancient DNA studies to reconstruct genealogies or population history (Izagirre and de la Rua 1999; Keyser-Tracqui et al. 2003) and to detect ancient sample contamination (Hummel et al. 2000). The HVl region of the mtDNA, because of its haploid and maternal mode of inheritance, gives information on the origin and genetic evolution of populations (Richards et al. 2000; Yao, Kong et al. 2002). The aim of this study is to determine the genetic affiliation of the Scytho-Siberian skeleton by analyzing these two genetic markers.

Materials and Methods

Source of Ancient Human DNA. During the summer of 1995 a Belgian and Russian scientific team excavated several Scytho-Siberian kurgans (stone tumuli) at the source of the Kizil River (Altai Republic). This site constitutes a necropolis of about 30 kurgans, at an altitude of 2180 m. The climatic conditions at this altitude allowed the formation of permafrost, which increases the chances of recovering frozen tombs with well-preserved human material (Burger et al. 1999). One of the kurgans (dated to the 4th-2nd century B.C. by carbon-14 dating of samples from beams of the burial chamber) contained a wooden burial chamber with a human adult skeleton associated with the traditional Scytho-Siberian burial gifts, and the remains of a horse (Bourgeois et al. 2000). Anthropomorphic analysis (Orban and Polet 1995) suggested that this skeleton (95-KBI-52) belonged to a mature adult male (35-45 years old) who exhibited characteristic lesions related to horse riding. Only a part of the right femur was used for DNA analyses.

Ancient DNA Extraction. A part of the right femur was abraded (to a depth of 4 mm) with a sanding machine (Dremel) to remove possible contamination from the bone surface. The sample was ground to a fine powder with a drill fitted with a surgical trepan.

Three independent DNA extractions were made according to the method of FiIy et al. (1998). For each extraction, 2 g of bone powder was dissolved in an extraction buffer [5 mM EDTA, 2% SDS, 10 mM Tris HCl (pH 8.0), 0.3 M sodium acetate, l mg/ml proteinase K] and incubated for 16 hr at 56°C. After an organic extraction (phenol/chloroform/isoamyl alcohol, 25:24:1, v/v/v), the aqueous phase was purified using the CleanMix Kit (Talent, France), a method based on the large DNA affinity for the silica in the presence of guanidium thiocyanate, and concentrated to 40 µl using Microcon-30 filters (Millipore, France).

Autosomal STR Analysis. Amplifications were performed with the Amp-FlSTR Profiler Plus Kit (PE Applied Biosystems, France). Nine STR loci (D3S1358, VWA, FGA, D8S1179, D21S11, D18S51, D5S818, D13S317, D7S820) and the amelogenin locus were simultaneously amplified. Each amplification was carried out in 10 µl of a reaction mixture containing 3.82 µl PCR reaction mix, 2 µl primer set, 0.182 µl AmpliTaq Gold, and 1-4 µl of the DNA extract. Cycling parameters were 94°C for 11 min, followed by 37 cycles of 94°C for l min, 59°C for l min, and 72°C for l min, and a final delay of 45 min at 60°C.

The PCR products were analyzed on an ABl Prism 3100 automatic sequencer (PE Applied Biosystems).

Mitochondrial DNA Analysis. Mitochondrial analyses were performed on the hypervariable region 1 of the mtDNA control region. This region was divided into two subregions (a and b), amplified with the primers L15989/H16239 (5'CCCAAAGCTAAGATTCTAAT-3'/5'-TGGCTTTGGAGTTGCAGTTG-3') and L16190/H16410 (5'-CCCCATGCTTACAAGCAAGT-3'/5'-GAGGATGGTGGTCAAGGGAC-3'), respectively. For subregion a, we also used the primer Hl6167 (5'-GGGTTTGATGTGGATTGGG-3') to resolve amplification problems linked to the polycytosine region located between nucleotide positions 16184 and 16193 (Szibor and Michael 1999).

DNA amplifications were performed in 50 ?? of a reaction mixture containing 2-6 µl of the DNA extracts, 10 mM Tris HCl (pH 8.3), 50 inM KCl, l .5 mM MgCl^sub 2^, l mg/ml BSA, 200 µl of each dNTP, 0.25 µM of each primer, and 2 U of Taq Gold Star (Eurogentec). Cycling parameters were 94°C for 10 min, followed by 38 cycles of 94°C for 30 sec, 48°C for 30 sec for HVIa or 30 sec at 51°C for HVIb, 72°C for 45 sec, and 72°C for 5 min.

Amplification products were checked on a 1% agarose gel and purified with Microcon PCR filters (Millipore). Sequence reactions were performed on each strand by means of the ABI Prism BigDye Terminator cycle sequencing Ready Reaction Kit (PE Applied Biosystems), according to the manufacturer's conditions. The sequence reaction products were analyzed on the ABl Prism 3100 automatic sequencer (PE Applied Biosystems).

Comparative Analysis. The nucleotide sequence obtained from skeleton 95-KBI-52 was compared with the mtDNA sequences of 8534 individuals gathered from the DDBJ/EMBL/GenBank international nucleotide sequence database and literature. Part of these sequences is included in the database compiled by Richards et al. (2000), which contains 2804 modern European, 1088 modern Near Eastern, and 208 modern northern Caucasian mtDNA sequences and is available at http://www.stats.ox.ac.uk/~macauley/founder2000/index.html. The other sequences constitute a second database with 323 modern northern Amerindian samples, 3710 modern Asian samples, and 401 modern west European samples (Table 1). All these sequences were aligned using the BioEdit 5.0.9 program (Hall 1999).

To determine the distribution of the 95-KBI-52 mtDNA haplotype in the reference populations, we used the Blast 2.0 program (available at http://www .ncbi.nlm.nih.gov). The haplogroup motifs reported by Richards et al. (2000) were used for haplogroup assignment, and the frequency of the 95-KBI-52 haplogroup was determined in each of the reference populations to assess the genetic affiliation, by maternal lineage, of this ancient subject.

Moreover, to complete this approach, we constructed a haplotype network (Bandelt et al. 1999) to depict the relationship between the Kizil haplotype and close haplotypes selected from among the 8534 reference haplotypes used in this study. We included only the haplotypes less than 5 mutations removed from the Kizil skeleton haplotype in the analysis. We used the median-joining method in the Network 3.1[beta] program (available at http://www.fluxus-engineering.com) to construct the network.



To investigate the population affinities of the Kizil skeleton, we used an assignment method to determine the population(s) that the STR profile obtained from the Kizil skeleton is most likely to occur in. We chose the frequency method, first presented by Paetkau et al. (1995), because, in using the allelic frequencies directly, it allowed access to a larger number of populations. The procedure was previously described by Cornuet et al. (1999). We used the allelic frequencies (corresponding to the STR loci analyzed in this study) from 22 modern populations, representing 4687 individuals, to perform the assignment method. The populations that we used included 2 eastern European populations (402 Russians and 455 Poles) (Pawlowski and Maciejewska 2000; Wolanska-Nowak et al. 2001; Kornienko et al. 2002), 2 central European populations (194 Austrians and 155 Bavarians) (Neuhuber et al. 1999; Anslinger et al. 2001), 3 southern European populations (115 Galicians, 110 northern Portuguese, and 143 Greeks) (Gusmao et al. 2000; Sanchez-Diz et al. 2002), 2 Middle Eastern populations (140 Egyptians and 310 Turks) (Klintschar et al. 1999; Asicioglu et al. 2002), 9 eastern Asian populations (178 Vietnamese, 300 Thai, 379 Koreans, 531 Chinese from Hong Kong, 198 Chinese from Beijing, 87 Chinese from Macao, and 300 Chinese from northeast China) (Han et al. 2000; Rerkammuaychoke et al. 2001; Wong et al. 2001; Law et al. 2002; Fung et al. 2001; Gusmao et al. 2001; Shimada et al. 2002; Wang et al. 2003), and 5 eastern Indian populations (65 Colla, 110 Brahmins, 103 Kayastha, 105 Meiti, and 101 Muslims) (Reddy et al.2001;Duttaetal. 2000).

Prevention of Contamination and Authentication. During all steps of sample preparation (abrasion, extractions, amplifications), we took all necessary precautions to guard against contamination from modern DNA. The workers who handled the samples always wore gloves, face masks, and lab coats, and the laboratory equipment (pipettes, tubes, filter tips, lab coats) was sterilized by exposing it to long ultraviolet light exposure. Laboratory rooms for DNA extraction, amplification, and post-PCR analysis were strictly separated. To detect possible contamination by exogenous modern DNA, we used extraction and amplification blanks as negative controls. Genotypes and mitocho
ndrial haplotypes of all persons involved in processing samples were determined and compared to the results obtained from the ancient bone sample. Moreover, DNA was extracted from the right femur at least three times, at different times; at least three amplifications were made from each extract to assess the reproducibility and the authenticity of the results.

Results

STR Analysis. The genetic typing by megaplex amplifications of the DNA extracted from the Kizil skeleton was not fully successful because a complete allelic profile could not be established (Table 2). Indeed, to reduce ancient DNA STR genotyping errors and to ensure the reliability of the results, we considered as authentic only the amplified products from each locus that were reproducible in at least five different amplification reactions (Schmerer et al. 1999). This strategy allowed identification of artifact alleles and false homozygosity resulting from sporadic contamination or amplification artifacts (such as shadow bands and allelic dropout) caused by the lack of DNA quality and/or quantity (Schmerer et al. 1999; Hummel et al. 2000). Consequently, the consensus profile determined after combination of different amplification results was incomplete (Table 2). For seven STR loci (VWA, FGA, D8S1179, D21S11, D18S51, D5S818, and D13S317) the obtained alleles are reproducible in five different amplification reactions and can be considered authentic (Schmerer et al. 1999), but for the other two STR loci (D7S820 and D3S1358) the degree of reproducibility of results and/or the rate of amplification success (44% for locus D7S820) were not sufficient to provide a reliable profile. For locus D7S820 this could be the consequence of the large size of the amplification products (256-292 bp) (Hummel et al. 2000), and for locus D3S1358 it was not possible to assess with certainty the homozygous or heterozygous nature of the genotype obtained.

The amplification results of the amelogenin locus were fully reproducible and consistent with the morphological sexual determination that 95-KBI-52 was a male.

The assignment method was performed from only the allelic frequencies of the seven STR loci considered in the consensus genotype. The probability of observing an individual with the Kizil skeleton STR profile was the highest in the two eastern European populations (Russia and Poland). Indeed, the likelihood that the Kizil skeleton STR profile occurred in these two populations was 10 times higher than in other European populations, 100 times higher than in eastern Asian populations, and about 100,000 times higher than in Indian populations.

Mitochondrial DNA Analysis. A 338-bp fragment (nucleotide position 16047 to 16385 of the CRS) of the mtDNA HVl region was sequenced and confirmed on both strands. This sequence (Table 3) was compared to the Cambridge Reference Sequence (Anderson et al. 1981), and there were seven variable nucleotide positions, represented by six transitions and one transversion. The mutations at nucleotide position 16147 C [arrow right] A, 16172 T [arrow right] C, 16223 C [arrow right] T, 16248 C [arrow right] T, and 16355 C [arrow right] T correspond to substitutions characteristic of the Eurasian haplogroup NIa (Richards et al. 2000). The haplotype comparison with the mtDNA sequences of 8534 individuals showed that this sequence was not found in any other population.

Discussion

In this study we have shown that the DNA recovered from the Scytho-Siberian skeleton from Kizil is relatively well preserved because a genetic profile and a mitochondrial haplotype could be determined. The positive results can be attributed to the good preservation of the skeleton, which was buried in a cold environment (the frozen grave being in direct contact with the permafrost); this environment was particularly favorable to the preservation of the DNA molecule (Burger et al. 1999; Leonard et al. 2000). The reproducibility of the results, the agreement between the morphological and molecular sex (Meyer et al. 2000), and the difference between the genetic profiles and mitochondrial haplotypes of the ancient skeleton and those of the technicians and archeologists (Table 4 and 5) are proof of the authenticity of the DNA samples studied.

To obtain the HVl sequence from the Kizil skeleton, we had to perform several amplifications from three independent extractions to resolve some ambiguities. The thymine (T) to cytosine (C) transition at nucleotide position 16189 led to the formation of a polycytosine region containing 10 cytosines. This polycytosine region, associated with the degraded nature of the DNA, did not allow the fixation of the primer H16239 and the subsequent amplification of the heavy strand of subregion a, including the putative mutation at nucleotide position 16147. To resolve this problem, we were able to read the heavy strand correctly by making use of the design of the primer H16167, which annealed before the polycytosine region. Thus we were able to confirm the C to A transversion at nucleotide position 16147 for both strands.

By using the haplogroup motifs described by Richards et al. (2000), we calculated the frequency of haplogroup NIa in the reference populations used in this study. The current frequency distribution of haplogroup NIa in the reference populations with this haplogroup is indicated in Table 6. The NIa haplogroup was not observed among the native American, east Asian, Siberian, Central Asian, and western European populations. The geographic distribution of haplogroup NIa is restricted to regions neighboring the Eurasian steppe zone. Its frequency is very low, less than 1.5% (Table 6), in the populations located in the western and southwestern areas of the Eurasian steppe. Haplogroup NIa is, however, more frequent in the populations of the southeastern region of the Eurasian steppe, as in Iran (but only 12 individuals were studied) and southeastern India (Karnataka and Andhra Pradesh territories). More precisely, in India haplogroup NIa is absent from the Dravidic-speaking population and is present in only five lndo-Aryan-speaking individuals, four of whom belonged to the Havik group, an upper Brahman caste (Mountain et al. 1995).

The HVl sequence from the Kizil skeleton belongs to haplogroup NIa, which is present only in Indian, Middle Eastern, and eastern European populations (Table 6) and therefore would be linked to European populations. This result is also supported by the haplotype network, which shows the relationship between the ancient Scytho-Siberian haplotype and haplotypes belonging to three Indian, Middle Eastern, and eastern European white populations (Figure 1). This is consistent with the hypothetical distribution of European populations in Central Asia during the 1st millennium B.C. (Lebedynsky 2001) and the results of previous mtDNA studies from the Scytho-Siberian population (Voevoda et al. 2000; Clisson et al. 2002). Indeed, the analyses by Clisson et al. (2002) and Voevoda et al. (2000) of the mtDNA HVl region from two and three Scytho-Siberian skeletons, respectively, showed that they belonged to five different maternal lineages affiliated with five different haplogroup characteristics of European and Asian populations. Besides, none of the five Scytho-Siberian mtDNA sequences previously published belong to haplogroup NIa, with which the Kizil skeleton is affiliated. This indicates that 2500 years ago the Scytho-Siberian population in Altai was of mixed Eurasian origin. The fact that the Kizil skeleton belongs to haplogroup NIa attests to the presence in Altai 2500 years ago of an individual linked by maternal lineage to a European population. The belonging of the six Scytho-Siberian skeletons to six different mitochondrial haplogroups could suggest, with respect to the weak number of Scytho-Siberian mtDNA samples available, a relatively important diversity of the Scytho-Siberian mitochondrial gene pool.

Moreover, haplogroup NIa was found with a relatively high frequency (8.3%) only in the Iranian and Indian Havik populations (Table 6). The ancient Scytho-Siberian skeleton could thus be linked to European populations living in the neighboring regions of this part of Central Asia, that is, the Indian and Iranian regions. The relationships, 3000-4000 years ago, between the populations of these three geographic areas (Iran, India, and Central Asia) are attested to by archeological records (Bokovenko 1994; Dvornichenko and Fedorov-Davydov 1994; Francfort et al. 2000; Lebedynsky 2001) and recent genetic studies on Y-chromosome markers and mtDNA (Quintana-Murci et al. 2001; Bamshad et al. 2001 ; Roychoudhury et al. 2001 ; Zerjal et al. 2002).

The assignment method based on the analysis of the allelic frequencies led to results that confirm the results of the mtDNA analysis, showing the higher affinity of the Kizil skeleton STR profile to European populations and notably to eastern European populations rather than Asian populations. The weak likelihood that the Kizil STR profile occurs in modern Indian populations could be explained by the practice, by Indian populations during the last 3000 years, of strict caste endogamy, which favors genetic drift (Bamshad et al. 2001; Majumder 2001). We cannot exclude this cultural practice as one of the causes of the high frequency of haplogroup NIa in the Indian Havik population.

The absence of the Eurasian haplogroup N1 a in the 490 modern individuals of Central Asia (Shields et al. 1993; Kolman et al. 1996; Comas et al. 1998; Derenko et al. 2000; Yao et al. 2000; Yao, Nie et al. 2002) suggests changes in the genetic structure of Central Asian populations, probably as a result of Asian population movements to the west during the past 2500 years.

We conclude that our analysis of genetic data obtained from a skeleton recovered in a Scytho-Siberian kurgan (2500 years old) links this ancient skeleton to several European populations that live in the neighboring region of Central Asia and shows that the Scytho-Siberian population contained a European component (Voevoda et al. 2000; Clisson et al. 2002). Future molecular genetic studies of ancient nomadic populations will be essential to appreciating the history of human migration in this area.

Acknowledgments This research was supported by a grant from the French Department of Research. We express our sincere thanks to Rosine Orban and Caroline Polet (Laboratory of Anthropology and Prehistory of the Royal Belgian Institute of Natural Sciences) for their assistance and to all the archeologists and anthropologists for the quality of the excavations.

Received 13 March 2003; revision received 10 july 2003.

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FRANCOIS-X. RICAUT,1,2 C. KEYSER-TRACQUI,1 J. BOURGEOIS,1 E. CRUBEZY,2 AND B. LUDES1,2

1 Laboratoire d'Anthropologie Moleculaire, Institut de Medecine Legale, I I rue Huinann, 67085 Strasbourg Cedex, France 67085.

2 Anlhropobiologie, Universite Paul Sabalicr, CNRS, UMR 8555. Toulouse, France 31000.

3 Department of Archeology, Ghent University, Ghenl, Belgium B-9000.

Copyright Wayne State University Press Feb 2004
Provided by ProQuest Information and Learning Company. All rights Reserved

[Vojvoda]

Originally Posted by Zrinski

Alans are sub-ethnos of old Iranic tribes. Namely the Sarmatians and Schytians.

Hello our Ossetian brothers.

Was it you who posted a map of Serbs & Croat tribes in Poland before their arrival to the Balkans?

[Zrinski]

Originally Posted by Vojvoda

Was it you who posted a map of Serbs & Croat tribes in Poland before their arrival to the Balkans?

You mean this? ->

[Lisowczyk]

So is it a possible to assume that the Poles today are ancestors of Sarmatians/Alans, BialoChorwaci(White Croats), and the serboi tribes who settled in western Poland?If so isn't it fair to say that Slavs descend from Sarmatian tribes since the Croats and Serbs were identified as Sarmatian tribes. In the 5th century BC Herodotus called the Slavs of antiquity "Scythian farmers".

[Menydh]

Interesting subject. Rusalka had some information on Sarmatians. Where is she when you need her?

[Zrinski]

I'd say not just Poles but all Slavs. I think there is more than enough, not just indication, but even archeological and historical proof to claim Balto-Slavic tribes are actually of Iranic stock. This can be proven even linguistically as Balto-Slavic languages are far closest of all languages(besides Iranic) to Sanskrt....old language of the Aryans(Iranic tribes).

[Milesian]

Interesting maps posted by TseBbe. - Historical maps of europe
If accurate, they seem to suggest that sometime between 600AD and 700AD, both Serbs and Croats moved south to their present geograpahical positions.

Looking back at the early maps, Iberia and Albania seem to have shifted quite a distance too. Being originally located in Azerbajan and Iran respectively

[Banat]

Originally Posted by Lisowczyk

So is it a possible to assume that the Poles today are ancestors of Sarmatians/Alans, BialoChorwaci(White Croats), and the serboi tribes who settled in western Poland?If so isn't it fair to say that Slavs descend from Sarmatian tribes since the Croats and Serbs were identified as Sarmatian tribes. In the 5th century BC Herodotus called the Slavs of antiquity "Scythian farmers".

It is another topic, but some think that all those Illyrians, Thracians, Sarmatians and others were only names for Slavic peoples. There is much controversy on that though, because some describe them as different peoples, some as similar with similar languages.

I know that some source (not from antic, though) describes Sarmatians as "Sirbi", some connect Sarmatians with Illyrians, and some Nestor of Russian areas in 11th century (if I'm not wrong) describes Illyrians as Balkan Slavs. It is however interesting how "the second most numerous people on Earth, after the Indians", the Thracians, completely vanished in less than a century, when Slavs as most numerous group in Europe appeared.

Considering Poles, I heard there was a lengend of brothers Lech (Leh) and Czech (Cheh), out of whom the peoples of Czechs and Lechs developed. In Serbian medieval sources, Poles were called Lechs (Leđani), until recent times.