MTDNA H2a1 ENCYCLOPEDIA MtDNA H2a1e MARC-OLIVIER RONDU P A RIS 202 6 Mitochondrial DNA H2a1 Phylogenetic tree. Maár et al. (2021) The MtDNA H2a1 haplogroup diverged from MtDNA H2 haplogroup (at the transition np 951; 163541) during Late Neolithic2 in Caucasus area and entered the steppe mating networks from North Caucasus farmers but then became widespread in steppe populations. The MtDNA H2a1 haplogroup reached Central Europe during the Yamnaya expansion and Central Asia during Early Bronze Age period. It is associated to the Indo-European dispersion. Expansion of MtDNA H2a1 during Late Neolithic and Early Bronze Age. MtDNA H2a1e diverged from MtDNA H2a1 circa 5200 years ago (Behar et al.2012) during the Late Neolithic period and emerged in elite cemetery in Samara River (Russia). During Early Bronze Age, it spread Westwards, north of the Carpathian range, and divided in three main roads, one leading to Scandinavia, one towards British Islands and one leading to the eastern Iberian coast by the Rhine-Rhone corridor. Cover picture: a vew of the Samara River (Russia) 1 2 Roostalu et al. 2007 7,713.4 ± 1,754.3 BP, Behar et al. 2012 IND. N° ID/ SEQUENCE GENDER MtDNA YDNA AGE DATATION LOCATION CULTURE BIBLIO. Utevka V, Samara River, Russia Samara Narasimhan et al. (2019) 11 I7489/58, Utevka V, Kurg. 1, Grave 1 F H2a1e 3326-2926 calBCE (3126) 16 I26780; (Krivyansky-9, kurgan 4, unidentified burial) F H2a1e 2906-2702 calBCE (2804) Krivyansky-9 (Rostov Oblast, Oktyabrsky District, Krivyanskaya Village) Russia, Don Early Yamnaya Lazaridis et al. (2024) 19 ODK1 Odessa Kurgan, burial 10 (Library ID S11850.E1.L2 / I11850) F H2a1e1 2874-2630 calBCE Odessa Kurgan 10, Ukraine Possibly of Catacomb archaeological complex Nikitin, A.G. et al. (2025) M H2a1e1 2866-2475 calBC (2670) Samborzec 1, Poland Corded Ware F H2a1e 2458-2206 calBCE Landau an der Isar, Germany Bell Beaker M H2a1e1a 2196 – 2031 calBCE NM 460-64, 2 Lillevasby, Zealand, Denmark Late Neolithic F H2a1e1 Isle of Man, Rushen, Strandhall Bronze Age 24 31 32 poz381, Samborzec 1, grave 21 RISE562/ Skel. RISE562, I5020; F0228, obj. 136/92 = gr.9 CGG_2_107464/ UCIAMS282674; National Museum registration NM 460-64, 2 34 I8582, 1983-65 40 K014; Keenoge 14 41 CGG106513/ UCIAMS272876 F H2a1e1a 47 PIR3116B M H2a1e 49 Maturus R1b1a1b1a1 ~30-40 2195-1973 calBCE 2023 – 1775 calBCE H2a1e1b TafVI-10 R1b1a1a2 KMR015/ 1571/ Grave 40 M H2a1e I2a2b 73 I27018, Burial 19 M H2a1e1 N1a1a1a1a2a1c1 88 CE-CP_31/ CPK020 M H2a1e1a P1 89 I29526.TW (tooth) F H2a1e1a 90 I29524.TW (tooth) F H2a1e1a 96 BAL003/ INVMG 1982.083 F H2a1e 97 Ind 13913:85:1 / BRC061 F H2a1e1a 101 Antro ID AP-AL1, Mol ID 2273 F H2a1e 106 GOG35 (Pit N°50) F H2a1e1a 8-10 F H2a1e1a Child F H2a1e1a 123 134 SK2937 / ST2937/ Layer 11 Skl ID 2256; burial 70; ID 906 135 PCA 0377 F H2a1e1a 142 ST0146/ SK0146/ Layer 2 M H2a1e1a1 Senilis Late Neolithic McColl, H. et al. (2024) 3790 +/-30 BP Rec de Ligno, Le Pirou, Valros Bronze Age Brunel et al. (2020) 600-300 CalBC 200 BCE – 100 CE 4th – 5th century CE 419-538 c CE 460-540 CE 650-780 cal. AD I1 Adult Patterson, N. et al. (2022) Pending Cassidy et al. (2017) Gerlev STR. Vej 10 Individual 1 MFG 114/97; Burial Mound, stone cist, Denmark 200 BCE 500 CE 30-40 McColl, H. et al. (2024) 2009- 1778 BCE H2a1e1a 57 Keenoge, Co. Meath. Ireland Juras et al. (2021) Allentoft et al. (2015) / Olalde et al. (2018) 668-874 CE 10th- 11th c. AD 940-1170 AD 1000-1200 AD 18th – 19th c. AD Taforalt, Grotte des Pigeons, Morocco Kamarakhevi Necropolis, Mtskheta district, Georgia Kipchakovo, Russia Iberomaurusian Belaya Pyany Bor Rym et al. (2016) Skourtanioti, E. et al. (2025) Gyuris, Balázs et al. (2025) Celje – Celeiapark, Slovenia Late Roman Vyas et al. (2025) Tarragona, Cambrils, Els Masos Late Roman Carrión, P. et al. (2024) Picts Morez, A. (2023) GermanicThuringian Gretzinger, J., et al. (2025) Langobard Frankish Coia, V. (2022) Balintore, Scotland Brücken, Mansfelder Land, Saxony-Anhalt, Germany Appiano San Paolo Castelvecchio/Eppan St. Paul Altenburg Gandía, Sanxo Llop (Valencia, Spain). Buried with probable father GOG34 Groenmarkt-2, Sint-Truiden, Belgium rural parish, Cherry Hinton, Cambridgeshire Santok, Lusz Voidvoship, Poland Groenmarkt-1; Sint-Truiden; Belgium Late Visigothic Late Medieval Oteo-Garcia G. et al. (2023) / (2024) Beneker, O. et al. (2025) Late Medieval Hui, R. (2023) Late Medieval Stolarek, I (2023) Late Medieval Beneker, O. et al. (2025) 11 UTEVKA V, KURGAN 5, SAMARA OBLAST, RUSSIA. FEMALE, MtDNA H2a1e In comparison with its predecessors, the Early Bronze Age Yamnaya culture developed in the Samara region (Volga), has the greatest similarity with the Khvalynsk culture. The origin of the new era is determined in connection with the origin and development of the Maikop culture of the Ciscaucasia. Some of the oldest radiocarbon dates from Yamnaya culture graves have been obtained from graves in the Samara oblast. The area around Utevka has more Bronze Age kurgan cemeteries than any other part of the Samara River valley. The sample published here is not among the oldest dates but adds to the body of Yamnaya individuals from this region. Based on its genetic affinities, we designate this individual as having the Western_Steppe_EMBA (Early to Middle Bronze Age) analysis label and Yamnaya_Samara split label (Vagheesh M. Narasimhan et al. 2019). Utevka V, 1/1 (I7489): Context date of 3326-2926 cal BCE (4430±25 BP, PSUAMS-5790); The Utevka V cemetery, located northeast of the modern village, was excavated by P. Kuznetsov in 1995. The kurgan 5, containing a single grave, appears to be associated to the Yamnaya culture. In the grave were a female 30 to 35 years old and a child covered by red ochre. The MtDNA H2a1 was sampled from the female adult. Kuznetsov P.F. (2010), PROBLEMS OF STUDYING THE EARLY AND MIDDLE PERIODS OF THE BRONZE AGE OF THE SAMARA VOLGA REGION, 40 years of the Middle Volga archaeological expedition, Kuznetsov P.F., 2010 / П.Ф.Кузнецов (2010), ПРОБЛЕМЫ ИЗУЧЕНИЯ РАННЕГО И СРЕДНЕГО ПЕРИОДОВ БРОНЗОВОГО ВЕКА САМАРСКОГО ПОВОЛЖЬЯ, 40 лет Средневолжской археологической экспедиции, Кузнецов П.Ф., 2010 Narasimhan, V. M. et al. (2019). The formation of human populations in South and Central Asia, Science 365, eaat7487 DOI: 10.1126/science.aat7487 Vagheesh M. Narasimhan et al. (2019), The formation of human populations in South and Central Asia. Science, 365, 6457. eaat7487(2019). DOI:10.1126/science.aat7487 Figure 11: Plan of Utevka kurgan 5 (credit P.F. Kuznetsov) Figure 12: Utevka kurgan 5. Female individual with Child (credit P.F. Kuznetsov) 16 KRIVYANSKIY-9, KURGAN 4, UNKNOWN BURIAL. FEMALE, MtDNA H2a1e Kurgan 4 is in the southern part of the Kurgan group. The height of the mound from the buried soil is 2.0 m (the maximum height above modern surface was 1.45 m), the diameter is 40 m. The visually definable boundaries of the mound are increased due to erosion and ploughing. 27 burials were recorded in kurgan 4. The earliest is grave 4: 11 (not sampled), dug in the EBA in the original ground surface before the construction of the kurgan embankment. The original mound was enlarged twice, associated with burials of the Early and Middle Bronze Age. The earliest phase of Kurgan 4 was built in the EBA over four graves: 18 (1 male),19 (2 males), 20 (3 males & 1 female), & 21 (5 males), placed on a small hill with a diameter of 10 m. These four graves contained 12 individuals: 11 males, all but one belonging to the I2a Y-haplogroup; and 1 female belonging to mt-haplogroup U5a2b. All were genetically unrelated within 3 degrees. The reason why males of various ages died and were buried together in group graves is not obvious. Perhaps an epidemic disease killed them. This small EBA cemetery was covered by the original kurgan 4 mound. The mound was enlarged by the addition of a second layer of the embankment in the MBA over the Catacomb-culture grave 27. After the completion of the second embankment, strong erosion of the Kadamovka tributaries occurred on the site, marked by clay-sand layers in parts of the mound. The third expansion of the Kurgan 4 mound was connected with a group of MBA Catacomb burials. Lazaridis, I., Patterson, N., Anthony, D. et al. The genetic origin of the Indo-Europeans. Nature 639, 132–142 (2025). https://doi.org/10.1038/s41586-024-08531-5 Figure: Schematic preliminary plan of Kurgan 1-7 at Krivyanskiy-9. (Image Lazaridis et al. 2024) 19 ODESSA «Слободка-Романовка» KURGAN, BURIAL 10. FEMALE, MtDNA H2a1E1 The Odesa Kurgan (46.488, 30.721) was studied in 1912-13 by M. F. Shkadyshko and A. V. Dobrovolsky. A detailed analysis of kurgan’s stratigraphy was published by V. G. Zbenovich and A. M. Leskov in 1965 and is one of the most important stratigraphic sites in the northwestern Black Sea region, essential to clarify the historical and fluvial connections between the ancient cultures of this territory. Analysis of the stratigraphy of the Odesa kurgan shows the presence of the main and three additional embankments. The main mound was erected over Burial 11 located in the center of the mound. The deceased was buried on the mainland yellow clay in crouched position, on the right side, head to the east. The skull is thickly painted with red paint……Most likely, the other ancient burials were also covered by small embankments, particularly the central burial 11, the embankment of which did not exceed 0.7 m in height. It was at this height that the slab covering the later burial 10 and the stones of the inner cromlech with a diameter of 4.25 m were discovered. The slab and the cromlech were placed when still open low embankment, which could only have been built over burial 11, since burials 4, 5, 6, also located in the center of the mound, they lie much higher. Yamna burials located both in the center and along the kurgan’s circumference are probably associated with two subsequent embankments. Catacomb and Babyno burials, as well as burials of the Late Bronze Age period, were subsequently added into the formed mound of the kurgan. I11850 ODK1, Odesa kurgan, Burial 10, female, 2874-2630 calBCE (4150±25 BP, PSUAMS-11224,) Burial 10, possibly of Catacomb archaeological complex, was lowered into the main mound of the kurgan. The burial pit was covered by a stone slab. The interred was in a contracted position on the back with legs falling to the right side. A stone circle was arranged around the burial. V. A. Dergachev, Yamnaya culture of the Carpathian-Danube region Vol. I. Catalogue of the burials CEP USM КИШИНЭУ 2023 A. V. Dobrovolsky. Excavations of a burial mound in the Odessa suburb of Slobodka-Romanovka. ZOOID. v. XXXII, 1915, pp. 121144. Nikitin, A.G., Lazaridis, I., Patterson, N. et al. A genomic history of the North Pontic Region from the Neolithic to the Bronze Age. Nature (2025). https://doi.org/10.1038/s41586-024-08372-2 Toshchev, G. N. Yeshche raz o stratigrafii Odesskogo kurgana. in Kurgany v zonakh novostroyek Moldavii 175–182 (Shtiintsa , Kishinev, 1984). 24 SAMBORZEC 1, GRAVE 21, CORDED WARE, POLAND. MALE, H2a1e1 The site at Samborzec (Sandomierz, Świętokrzyskie Voivodeship, Poland), excavated in 1961-1966, locates on the border of loess upland, on the left bank of the Gorzyczanka River. It is at the southern periphery of a concentration of Late Neolithic grave sites that extends along the Vistula terrace from Sandomierz, via Złota, to Żuków and Samborzec. Four graves (19, 21, 22, 23) of the Corded Ware culture were discovered, a few meters apart of each other. The grave 21 was probably of the niche type. The burial pit had an oval shape. It was filled with patchy, light brown earth in the upper parts, and slightly darker and more uniform in the lower parts. The skeleton of a man aged maturus (poz381) was discovered at a depth of 80 cm from the surface of the earth. A human bone sample was conventionally radiocarbon dated to 4080±50 BP (Ki-7930) (Tunia, Włodarczak 2002). He was placed on his back, with his head and legs turned to the left side, oriented along the axis NE-SW, head to NE, face to E. The hands, slightly bent at the elbows, are placed on the pelvis. The legs of the individual were bent in a way typical for the Cracow-Sandomierz group with an angle in the hip about 90 degrees. Two ceramic vessels standards of the Cracow-Sandomierz group were found below his strongly curled legs. A small four-eared amphora (height 15.8 cm, spout diameter 11.2 cm, belly diameter 19.4 cm, bottom diameter 5 cm) with an undefined bottom and an arched outward rim preserved in its entirety. Decorated with seven bands of horizontal imprints of a left-handed rope and a band of arches, also imprinted with a rope. The ears are very wide with small holes (3-4 m in diameter). The outer surface is even, matte, in color orange-brown. Visible traces of wiping with panicles. The second vessel was an undecorated four-handed amphora with a slightly separated bottom (height 10 cm). It is lost today, only his drawing has been preserved. In addition, a bone awl made of bone (length 88 mm) was discovered in the filling of the cavity, above the skeleton. The characteristics of the funeral rite and artifacts indicate that the cemetery in Samborzec belong to the younger stage of Corded Ware culture in Lesser Poland. The Graves nos. 19, 21, and 22 belong perhaps to the youngest developing phase of the Cracow-Sandomierz group. Juras et al. (2021). Maternal genetic origin of the late and final Neolithic human populations from presentday Poland. https://doi.org/10.1002/ajpa.24372 Tunia, K., Włodarczak, P. (2002), Radiocarbon Results for the Corded Ware Culture from Southern Poland. Przegląd Archeologiczny 50, 45–55. Włodarczak, Piotr (1999), Cmentarzysko w Samborcu, woj. świętokrzyskie na tle innych znalezisk kultury ceramiki sznurowej z Wyżyny Sandomierskiej, Sprawozdania Archeologiczne, T. 51, 1999; PL ISNN 0081-3834 Polska Akademia Nauk. Instytut Archeologii i Etnologii, Kraków, 1999 / Sprawozdania Archeologiczne Vol. 51 (1999); The Cemetery at Samborzec, Świętokrzyskie Province, on the Background of Other Finds of the Corded Ware Culture on the Sandomierz Upland Figure 24: the Samborzec grave plan and inventory (credit Włodarczak, Piotr) 31 LANDAU AN DER ISAR (DINGOLFING-LANDAU), BAVARIA, GERMANY. 2458-2206 calBCE. BELL BEAKER GRAVE 9. FEMALE INDIVIDUAL MtDNA H2a1e “On a terrace above the Isar river (a tributary of the Danube) in the outskirts of Landau town, a Bell Beaker settlement and an adjacent cemetery were excavated in 1992. The graves were nine inhumations, of which two were double graves, and two were cremations. In 1981, a single, richly furnished, Bell Beaker grave was excavated ca 80 m further to the west. The graves in the cemetery were also rich, with items such as gold and copper objects. Six graves were arranged in a NW-SE line, while the remaining three were somewhat further to the west. One individual from grave 9 was sampled for the Rise project (RISE562). This somewhat disturbed grave contained the skeleton of an adult female, lying on her right side. The cranium was found by the pelvis, probably due to disturbance. In the grave were 12 bone buttons with V perforations, a copper awl, and two red painted beakers with horizontally zoned decoration. Unique for Bavaria, a thin rectangular gold sheet, ca 6x3 cm, with perforations was also found. At the feet were bones from half a young pig. Here, another four individuals were analysed, from graves 3, 4, 5 and 7. Grave 3 was a double grave, containing two children, one ca 4 years and one ca 5 years old81–83. The skeletons were lying in tucked “hocker position” and facing in opposite directions. Each child was accompanied by an undecorated vessel. Grave 4 contained the skeleton of an adult woman. She was lying in right hocker position with head to the SE. At her feet were two undecorated handled vessels. Grave 5 was of an adult female, lying in right hocker position with the head to the SE. At her feet was a single undecorated handled vessel. Grave 7 was of an older adult woman. The somewhat disturbed skeleton was lying in right hocker position with the head towards the SE. In the chest region were 8 V-drilled bone buttons, by the left lower arm was a copper dagger, and at the feet were three undecorated vessels. Sr isotopes on bone and tooth samples from six individuals from Landau were measured by Price et al (2004)84. They suggest the individual in grave 7 was born locally while all others were migrants. Excerpt from Olalde et al. (2018) “Besides the presumed burial mound of Trieching (Grave l with circle ditch), there is a confirmed Bell Beaker tumulus at Wallersdorf. It is the first proven tumulus for that period in Lower Bavaria. Grave l at Trieching is the oldest burial in the Dingolfing-Landau Region. Assemblies for making fire were found in this grave, in grave 1/96 at Aufhausen, and in grave 2/92 at Landau. These assemblies were composed of a piece of flint and a piece of Iron Pyrite. In this case the Iron Pyrite was gone, however, the flint was coloured by it. This type of "lighter" has been known since the Linear Pottery Culture. The only flint daggers recovered from Bell Beaker burials were found in grave A at Landau, within the tumulus at Wallersdorf, and in a Bell Beaker grave at Pörndorf-Widdersdorf. The flint material comes from Arnhofen. The materials that comprise the copper daggers from the graves at Trieching, Landau (1981) and 1992 (from a female grave), as well as that recently discovered at Aufhausen, have not as of yet been examined as to their origins. It is quite conceivable that thoroughly different regions of origin could be established. The fact that a dagger lay in a female grave is quite remarkable and seemed to be quite rare within the Bell Beaker culture. Three small, thin bracelets out of gold were found in Grave 9/92 at Landau and in a newly uncovered cremation burial at Aufhausen. They belong to the oldest artefacts made from gold in all Europe. There are now two 14C dates for the "gold grave" at Aufhausen, their mean value being 3860 ± 45 BP. The dating was made through the kind assistance of Dr Lan Lanting at the University of Groningen in Holland. Both bell beakers from grave 9/92 in Landau and the cremation grave at Aufhausen (the grave of a young man) present very similar decoration patterns. This could indicate a kindship relation between the woman buried in Landau and the man buried in Aufhausen. Gold foils are a type of hair ornament and seem to lie predominantly in female graves. Besides a recently discovered female grave with gold foil in Moravia (this body grave was discovered in 2001 in the proximity of Brno and still unpublished), there is another "gold grave" in Moravia, two in France, two in Spain, and now two in Niederbayem, Germany. These las t two are only about 10 km from one another. Future DNA testing of the skeletons may well confirm the kindship relations that have been speculated with up to now, based upon their identical grave goods including quite similar ornamental patterns on the beakers. Therefore, the district of Dingolfing-Landau would seem to be one of the most important centres of the Bell Beaker Period in Bavaria. Excerpt from Kreiner, L. (2001) “Just 1 m north of grave 6 was the almost 0.5 m deep grave pit of grave 9, at the upper level of which it was already clear that it had been disturbed or robbed. After the excavation, it was also apparent that the adult woman's upper body was completely disorganized. 20 Where the skull once lay, a copper awl with two pointed ends and a thin gold-electron-copper sheet (6.3 x 2.6 cm) were found, which was perforated three times on each of the sides of the skull and had eight tiny punched indentations on a rounded corner. Half of a pig21 and two decorated cups placed one inside the other with traces of red painting were found on the feet of the dead. 22 The cups date the grave to the middle stage (stage A2 according to V Heyd) of the Bell Beaker culture. Thus, grave A (1981), 1/92, 6/92 and 9/92 date to the middle period, the remaining graves appear to have been created in the same period, but somewhat later. The graves and the findings from two large settlement pits dating from the same period, which were uncovered about 120-190 m south of the graves, give the impression that a family group had lived and been buried here for two to three generations.” Excerpt from Kreiner, L. (2001). “The two vessel units from Landau an der Isar (HD-19835, 2433–2209) are in a maritime/epi-maritime context. These squat vessels (B 2.5) are also provided with narrow decorative bands that include the foot area. These are filled with horizontal lines (10) and bundles of lines (18) as well as diamond bands”. Excerpt from Großmann, R. (2016) MtDNA H2a1e The Mitochondrial DNA sub-haplogroup H2a1e probably diverged from the macro-haplogroup H2a1 in the Pontic Steppe or Volga area and slowly migrated towards central Europe until Landau an der Isar. The most ancient sequence has been retrieved from the Female individual buried in Utevka site V, Kurgan 1, grave 1 (I7489/ 58; 3326-2926 calBCE, Figure 4 N°9), Samara River, Russia (Narasimhan et al. 2019). It was later sequenced in the site Krivyansky-9, Kurgan 4 (Rostov Oblast, Oktyabrsky District, Krivyanskaya Village, Russia) with the female individual MtDNA H2a1e (I26780; 2906-2702 calBCE, Figure 4 N°13) and associated to the Early Bronze Age on Don dominated by the Yamnaya culture (Lazaridis et al. 2024). The migration towards west continued and the sub-group MtDNA H2a1e1 was sequenced from an individual buried in the site of Samborzec 1 (poz381, grave 21, 28662475 calBC, Poland, Figure 4 N°19) dominated by the Corded Ware culture (Juras et al. (2021). From now on, the migration of MtDNA H2a1e carriers towards Northern (Denmark) and North-Western Europe (British Islands) accelerated, and in a second impulse, they followed the Rhone and the Iberian Peninsula to reach North Africa. Allentoft ME, et al. (2015), Population genomics of Bronze Age Eurasia. Nature. 2015;522:167–172. Bosch, Tobias Ludwig (2009) Archäologische Untersuchungen zur Frage von Sozialstrukturen in der Ostgruppe des Glockenbecherphänomens anhand des Fundgutes. PhD, Universität Regensburg Großmann 2016, GROßMANN R., Das dialektische Verhältnis von Schnurkeramik und Glockenbecher zwischen Rhein und Saale, Bonn, Rudolf Habelt, 2016, 279 p. (Universitätsforschungen zur prähistorischen Archäologie; 287 / Human Development in Landscapes; 8) Heyd, VM., Husty, L., & Kreiner, L. (2004). Siedlungen der Glockenbecherkultur in Süddeutschland und Mitteleuropa. Büchenbach Verl Dr Faustus Husty, L. (1992), Ein Gräberfeld der Glockenbecherkultur in Landau a. d. Isar. Das Archäologische Jahr in Bayern 1992 Juras et al. (2021). Maternal genetic origin of the late and final Neolithic human populations from present-day Poland. https://doi.org/10.1002/ajpa.24372 Kreiner, L. (2001), Die Graber der Glockenbecherkultur im Landkreis Dingolfing-Landau, Niederbayern, Preistoria Alpina, v. 37 (2001), pp. 337-363, Trento 2002 Lazaridis, I. et al. (2024), The Genetic https://doi.org/10.1101/2024.04.17.5895971 9 Origin of the Indo-Europeans, bioRxiv 2024.04.17.589597; doi: Narasimhan, V. M. et al. (2019). The formation of human populations in South and Central Asia, Science 365, eaat7487 DOI: 10.1126/science.aat7487 Olalde I. et al. (2018), The Beaker phenomenon and the genomic transformation of northwest Europe. Nature. 2018 Mar 8;555(7695):190-196. doi: 10.1038/nature25738. Epub 2018 Feb 21. Erratum in: Nature. 2018 Mar 21;555(7697):543. PMID: 29466337; PMCID: PMC5973796. Price, T. D., Knipper, C., Grupe, G. & Smrcka, V. (2004), Strontium Isotopes and Prehistoric Human Migration: The Bell Beaker Period in Central Europe. Eur. J. Archaeol. 7, 9–40 32 LILLE VASBY, KOBENHAVN, SMORUN, SENGELOSE PARISH, ZEALAND, DENMARK. 2196 – 2031 calBCE. CGG107464/ UCIAMS-282674. MALE INDIVIDUAL MtDNA H2a1e1a. Jordfæstegrave, literally a grave in earthing trench, is already in use in the Stone Age (approx. 12,800-4,000 BC). The dead were placed unburned in earthen graves, wooden coffin, or skins, in the ground or under burial mounds. The burial form has been very widespread in Denmark except for a period spanning Late Bronze Age and pre-Roman Iron Age dominated by cremation. The earthen graves are found in many different forms, from the simpler constructions during Hunter Stone Age to the earthen graves of the Iron Age and the Viking Age, where in certain cases the bodies are buried in wagon, boats, or wooden burial chambers. “Depositions of bones in pits in connection with flat graves from the Late Neolithic have only been found in southeastern Scandinavia. At the burial site at Egedal there were two pits with human bone fragments. In another two graves the bones had been stirred around and one of the deceased had been dismembered (Pedersen 1993). In Lille Vasby in Zealand three individual burials and a mass grave were found. Nearby depositions of bones appeared, showing traces of dismemberment (Liversage 1966; 1980, 24 ff.). At the burial site at Stockholmsgården in Scania there were two pits with human bones (Strömberg 1951, 11 ff.). Such bone depositions can also be seen in connection with stone cists and in megalithic tombs. The reasons for such treatment have been discussed, and ethnographic parallels have been drawn (Jensen 2001, 531; Kaul 1994; Liversage 1966). One common assumption is that the depositions should be viewed in connection with rituals and ancestor worship (Jensen 2001, 531). In the graves at Bulbrogård there were no traces of repeated burials, so it is not likely that the pit was associated with the clearance of earlier burials. Excerpt from Bican, Josefine (2012). Lille Vasby: “A small fully excavated cemetery. The burial ground was excavated in 1964 by D. Liversage. The burial ground was placed above what has been interpreted as a Late Neolithic ritual place, containing scattered human bones (Liversage 1980:24). The cemetery contains 19 inhumation graves of which six are dated to the Early Roman Iron Age; the remains are either undated or belong to the Late Neolithic (Sellevold et al. 1984: 85). Grave 16, skeleton 72 is radiocarbon dated to the Early Roman Iron Age (1-120 CE).” Excerpt from McColl, H. et al. (2024). “At Lille Vasby between Copenhagen and Roskilde one found, for example, such a collection of flat field graves. These were both regular individual burials, a communal grave and burials with scattered bones. Some of the bones, including five children's skulls, were laid to rest at the edge of a large, irregular burial. Two of the skulls were each in a small depression of the burial. In addition, vertebrae, ribs, finger and toe bones of both children and adults were found lying scattered in the burial. In some places there were piles of limb bones, and several of these had been broken over before the demolition. However, they were not broken lengthwise, as they would have been if it was the bone marrow that had been sought to be obtained. In some cases, however, it could be established that the bones had been cut while connective tissue was still attached to them. In some cases, the bodies have therefore been dismembered even before the skeletonization. Where the bones came from is not entirely clear. But in the large nearby mass grave, skeletal parts of three adults were found. However, the skulls and most of the limb bones were missing – they had clearly been removed from the grave. What remained were vertebrae, ribs and other small bones. Here it was clear that the movement of the bones had taken place after the bodies had been skeletonised. As opaque as the rituals that lay behind this kind of bone movement may appear, they seem to be best explained on the basis of the ancestor worship that took place throughout the Peasant Stone Age, and which required that the bones of the dead be split apart. Excerpt from Jørgen Jensen, P. (2012) Bican, Josefine. (2012). Bulkister og Bådgrav ved Bulbrogård – en senneolitisk gravplads ved vestbredden af Tissø. AArbøger for Nordisk Oldkyldighed og Historie. 7 – 42. Jørgen Jensen, P. (2012), De levende og https://danmarksoldtid.lex.dk/De_levende_og_de_d%C3%B8de_-_2400-2000_f.Kr. de døde (2400-2000 f.Kr.). Liversage, D. (1980), Material and Interpretation: The Archaeology of Sjællamd in the Early Roman Iron Age. The National Museum of Denmark: Copenhagen. McColl, H. et al. (2024), Steppe Ancestry in western Eurasia and the spread of the Germanic Languages, bioRxiv 2024.03.13.584607; doi: https://doi.org/10.1101/2024.03.13.584607 Sellevold, B. Hansen, U.L., & Jørgensen, J.B. 1984. Iron Age Man in Denmark. Prehistoric Man in Denmark, Vol III. Københaven: Nationalmuseet. 34 STRANDHALL, RUSHEN, ISLE OF MAN. 2195-1973 calBCE. FEMALE INDIVIDUAL MtDNA H2a1e1 The site at Strandhall (Manx National Heritage NMHER 57) is located on the A5 between Kentraugh and Balladoole. During agricultural work in 1983, farm workers had stumbled across a prehistoric burial cist. Larch Garrad, from the Manx Museum went to investigate and carried out two small-scale rescue excavations. The short cist is situated on a ‘low hill with a surprisingly wide view’ (Garrad, n.d.a) and was discovered when a plough displaced the capstone. A course of stones to the south of the cist may derive from a kerb, with a projected arc implying a mound 10m in diameter, though no mound was extant at the time of excavation. The cist was 1.05m long and 0.4m wide, oriented north-north-east/south-south-west. The cist was located within a cutting into the bed-rock. It had no floor slab and three of the sides were made of a single slab of slate with the fourth side constructed from two blocks. Several additional stone boulders (probably taken from the beach) were also used to build-up the height of the cist walls and support the sides. The site contains two sets of human remains. The cist itself contained an inhumation burial with approximatively 5% of the unburnt skeletal remains of a 10–13-year-old child (accession number 198365; Gamble 2017). The excavator considered the remains to be part of a crouched inhumation, but the finders had moved some of the bones prior to the excavation. The burial might have been disturbed by an animal as some of the teeth were not located near the skull. A thumbnail scraper, flint flakes, and shells from land snails (possibly intrusive) were found in the cist. A tooth from the child has been radiocarbon dated to 3685±28 BP (OxA-37603; 2195–1973 cal BCE). This burial is the only known case of the inhumation of a child from this period on the Isle of Man, although there are contemporaneous burials of the cremated remains of children. Beyond the projected kerb, roughly 2m east of the cist, and sharing the same alignment, was a stone-lined trench c. 0.75m wide and 7m long. A substantial amount of cremated bone (accession 1983-0201) was found at the south end. A sherd of an Early Bronze Age Cordoned Urn along with shells from limpets, winkles, and common garden snails (the latter possibly intrusive) lay within the trench (Garrad n.d.b.). The bone could derive from a single individual and a tooth suggests an age at death of c. 16–22 years, based on wear (though this is tenuous). A sample of long bone was radiocarbon dated to 3859±29 BP (OxA-36596; (2459–2206 cal BCE). Sample I8582 (female) derives from the left first maxillary molar of the 10–13-year-old child in the short cist (1983-65). The tooth has also yielded stable isotope readings of -20.65 for δ13C and 11.8 for δ15N. These results reflect diet before the age of 10 years and the nitrogen value suggests some consumption of marine foods (Ashley Coutu, pers. comm.). The site will be reported on fully in a monograph summarizing the work of the Round Mounds of the Isle of Man project. Excerpt from Patterson et al. (2021). Author of entry: Chris Fowler and Michelle Gamble Strandhall Farm is located on the A5 between Kentraugh and Balladoole in one of the farm fields. The site was found during agricultural work in 1983. The farm workers had stumbled across a prehistoric burial cist. Larch Garrad, from the Manx Museum went to investigate what had been found and carried out two small-scale rescue excavations. Garrad initially opened a trench large enough to investigate the cist. The cist was located within a cutting into the bed-rock. The cist had no floor slab and three of the sides were made of a single slab of slate with the fourth side constructed from two blocks. Several additional stone boulders (probably taken from the beach) were also used to build-up the height of the cist walls and support the sides. The cist itself contained and inhumation burial. The cist had filled with soil and Garrad suggests it might have been disturbed by an animal as some of the teeth were not located near the skull. Garrad also found a scraper and a flint flake in the cist. Near to the cist Garrad found a band of waterworn boulders located within a yellowish soil – which she interprets as the remains of a stone kerb that once surrounded the mound. She used the location and size of this spread of stones to estimate that the mound might have been about 10m in diameter. In addition to the burial in the cist a small deposit of cremated remains were found nearby. Garrad’s excavated the site on two separate occasions and the cremated remains appears during this second excavation and it is a little tricky to understand how these cremated remains relate to the mound. If we think that Garrad had accurately identified the kerb-edge of the mound then these cremated remains were found outside of the main area of the mound in what looks to have been a stone-lined trench nearby. Garrad suggests this stone setting interrupts the second cist containing the cremated remains. The cremation was found under a small stone slab that she interprets as a capstone. Garrad tells us she found a sherd of Collared Urn, a form of Bronze Age pottery, nearby in the stone setting potentially dating the feature to the Bronze Age. The relationship between the stone setting and this second cist is hard to work out, the stone setting may well be a potentially later drainage ditch. Garrad also notes that the fragments of the cremated skull had been placed on top of the rest of the cremated remains. This suggests that the body was cremated elsewhere, then bones were collected and brought to the site for burial with the mourners selecting specific fragments of the skull to place on top of the rest of the bones. Michelle examined the bones from Strandhall in the Manx Museum in October. The bones from the inhumation burial in the cist are badly eroded from the acidic local soil. Less than 5% of the skeleton remains. The teeth from this burial and the nature of the bones suggest that the person died around age 10-13 years old. It was not possible to work out their sex. The cremated bones appear to also belong to a relatively young person – aged around 16-22 years when they died. These bones are quite small and highly warped from the cremation process. Michelle could not identify the sex of this individual either but was able to suggest they were perhaps a small person. Within the cremated bone she also found a small amount of unburned animal bone and flint which appears to have been deposited with the cremation. It may have been that a cut of meat for example was buried with the cremated remains. Strandhall – the burial of two young people Author: Rachel Crellin and Michelle Gamble (2017) https://roundmounds.wordpress.com/2017/04/07/strandhall-the-burial-of-twoyoung-people/ Crellin, R. & Gamble, M. (2017), Strandhall – the burial of https://roundmounds.wordpress.com/2017/04/07/strandhall-the-burial-of-two-young-people/ two young people Gamble, M. 2017. Strandhall, Rushen. Human Osteology Report. Unpublished manuscript, Manx National Heritage. Garrad, L. n.d.a. Cist at Strandhall, Rushen. Unpublished manuscript, Manx Museum Library. Garrad, L. n.d.b. Excavation at Strandhall, Phase II. Unpublished manuscript, Manx Museum Library. Patterson, N., Isakov, M., Booth, T. et al. Large-scale migration into Britain during the Middle to Late Bronze Age. Nature (2021). https://doi.org/10.1038/s41586-021-04287-4 40 KEENOGE, MEATH, IRELAND. 2023 – 1775 calBCE. K014, Co. INDIVIDUAL MtDNA H2a1e1b (UNPUBLISHED) “An Early Bronze Age flat cemetery at Keenoge, Co. Meath, in use some time between 2280 BC and 1520 BC, consisted of six cist burials and eight-ten pit burials, which contained at least 26 individuals. At least seven of the graves contained pottery and one contained bronze and flint artefacts. The finds consisted of the remains of up to twelve complete or partially complete vessels, including nine complete bowl food vessels, one or two vase urns, one encrusted urn, a jet necklace of 40 beads, one bronze razor knife, one flint scraper and a number of other flints. The site was excavated between 1929 and 1936 by Dr Mahr, director of the National Museum from 1927 to 1939. Grave 14: pit burial This was a pit (PL XVI; Fig. 3) but there is no information on the form of the grave or its dimensions other than that it was lined with a few stones. The grave is recorded as containing a single crouched inhumation (1929:72b), but examin ation of the bone revealed that two inhumations, an adult male and a juvenile or adolescent, were present, as well as the partially cremated remains of a third and fourth individual, an adult male and an infant, indicated by a fragment of skull. As with grave 12, a primary burial may have been disturbed and a second inhumation and a cremation inserted. The crouched burial was associated with a bowl food vessel (Fig. 16, 1929:73), which was found near the head. This is an unusual hemispherical bowl with an almost pointed internally bevelled rim and a thick pronounced foot. The ware isvery thick. It is decorated with short vertical lines on the rim bevel and a series of horizontal arc-shaped impressions on the exterior separated by incised lines, with slanting incised lines on the loot. The vessel measures 9.6cm in height, 8.3cm in rim. Excerpt from Mount, C. (1997) Kassidy et al. (2017 unpublished Mount, C. (1997), Adolf Mahr’s excavations of an early Bronze Age cemetery at Keenoge, County Meath, Proceedings of the Royal Irish Academy, Vol. 97C, 1-68 (1997) 41 GERLEV, STRANDVEJ 10, SJEALAND, DENMARK. 2009- 1778 BCE. MFG 114/97, / INDIVIDUAL 1MtDNA H2a1e1a “The two samples come from the southernmost of two burial mounds called Iversbjargene, which were previously found close to the western shore of Roskilde Fiord. The remains of the mound were excavated after a so-called fishtail flint dagger (of Lomborg's type IV) was found on the site in the late 1990s. The dagger indicates that the burial mound belonged to the last phase of the Late Neolithic. The excavation revealed the last remains of the mound, which seems to have been surrounded by a stone frame. The mound contained an east-west oriented gallery grave covered by a stone package including several shells. The top stones of the cist were missing, but the side stones consisted of slender split stone slabs. Remains of a white limestone pavement were preserved in the floor. The internal dimensions of the cist were 250 x 100 centimetres. The unburned bones of at least three individuals were found scattered in the filling. Fragments of burnt bone, also found in the filling, together with a fragment of a serrated bronze sickle, may represent the remains of a Late Bronze Age cremation. The bones of individuals 1 and 2 were selected for study: Individual 1 was identified as a male between 25-35 years old (closer to 25 than 35). The bones were strong and large, and the presence of prominent muscle attachment markings on the knee and heel bones suggest a muscular physique. Mild dental calculus was observed. There was a lack of wisdom tooth development (agenesis) in the preserved mandible. Individual 2 was probably a male over 40 years old. His bones were not as robust as those of Individual 1, but still somewhat stronger than the unexamined Individual 3. Significant tooth loss (with signs of healing during the individual's lifetime) was observed in the preserved left half of the jaw. There was also evidence of osteoarthritis in the lumbar vertebrae and a finger joint. The third individual, who was not sampled, had only a fully preserved right humerus, a left kneecap (patella), a left talus (ankle) and a right navicular (foot). These bones were more delicate and smaller than those identified and sorted as belonging to Individuals 1 and 2, suggesting that they belonged to a third, possibly female, individual. The individual was adult, i.e. over 18 years of age, but was probably older (at least over 30 years of age) as the joint surfaces, where they could be observed, were not as sharp as in younger individuals. Based on the humerus, the stature is estimated to have been between 154 and 157 centimetres. A small stone cist was found just outside the stone frame of the mound. Nothing was found in it. and function remains uncertain. Excerpt from McColl, H. et al. (2024), supplementary files by Jens Winther Johannsen. McColl, H. et al. (2024), Steppe Ancestry in western Eurasia and the spread of the Germanic Languages, bioRxiv 2024.03.13.584607; doi: https://doi.org/10.1101/2024.03.13.584607 47 REC DE LIGNO, LE PIROU, VALROS, FRANCE. 3790 BP. PIR3116B. MALE INDIVIDUAL MtDNA H2a1e The burial feature Ens 11 is located west of the other archaeological structures. It is located in a pit orientated East-West (1,54mx 1,13mx 0,20m). The pit has yielded two adult individuals. This is a primary burial for the female individual 11A, lying in a flex position on right side, East-West oriented, facing west. The lower limbs have undergone changes, only the proximal part of the left femur remains in connection with the acetabulum. This disturbance corresponds to the area where a stone assemblage has been observed. Three blocks (two in limestone, one in basalt) appeared on the surface located near the eastern wall of the pit. They are arranged parallel with a dip towards the north. Under the southernmost slab, an alignment of three stones (sandstone and limestone) arranged east–west was isolated which rested on the bottom of the pit, probable indirect evidence of a structure. On the remains of the Female individual 11A, were deposited the bones of a second adult 11B aged between 20 and 29 years old. The long bones are found in bundle to the north of Individual 11A. The other elements of the skeleton (ribs, hip bones, vertebrae, etc.) are located to the northeast of the individual 11A and the craniofacial block on the spinal column of the skeleton in connection. The absence of preservation of anatomical connection and the arrangement in a bundle allow us to attest to a secondary deposition of these bones. However, the presence of different elements of small modules (phalanges, metacarpals, metatarsals) suggests the body decomposed in the pit. The grouping of bones in a secondary position on the remains of individual A and the fact that certain long bones (tibia, femur) are kept stacked in balance testifying to an effect of constraint would indicate that these remains were wedged between the body of the deceased and a wall having disappeared. The bones in the secondary position of individual B covering those of subject 11A in position suggest that the latter was also deposited with the skeletal remains, from a first burial, within a rigid container made of perishable material of formwork type with possibly a bottom. Brunel, S. et al. (2020). Ancient genomes from present-day France unveil 7,000 years of its demographic history. Proceedings of the National Academy of Sciences. 117. 201918034. 10.1073/pnas.1918034117. Y. Gleize · G. Loison · R. Marsac (2013), Gestion singulière des morts au sein d’un petit ensemble de tombes du Bronze ancien à Valros (Hérault) Bull. Mém. Soc. Anthropol. Paris (2013) 25:190-207 DOI 10.1007/s13219-012-0070-6 Figure: The Ens 11 burial pit (graphics F. Vinolas) in Gleize, Loison, Marsac (2013) fig.7 p. 199 57 KAMARAKHEVI NECROPOLIS, MTSKHETA DISTRICT, GEORGIA. 600-300 CalBC. KMR015/ 1571/ G RAVE 40, MALE MtDNA H2a1e Kamarakhevi (KMR) A) General Location and Chronology The Kamarakhevi cemetery (41.856617, 44.732793) is located to the south of Tsitsamuri village (Mtskheta district), on the plain along the left bank of the river Aragvi and its tributary the Great Kamarakhevi. The tombs discovered at Kamarakhevi are divided into three chronological groups: Group I (6th -5th centuries BCE), Group II (5th -4th centuries BCE), and Group III (4th -3rd centuries BCE). Five radiocarbon dates are in agreement with the archaeological date, while one earlier radiocarbon date (1385-1202 calBCE) indicates an earlier phase of cemetery use. B) Excavation history and description of burials Archaeological excavations were carried out in 1953 and 1976-77, led by T. Zhgarkava, A. Afakidze, and R. Ramishvili. The initial excavation in 1953 uncovered 39 tombs. Some tombs yielded large quantities of jewellery and pottery, while the majority of child burials were devoid of burial goods. The predominant burial type was pit burial, followed by cist grave. To date, more than 100 tombs have been excavated at Kamarakhevi. Burial customs across the cemetery appear relatively uniform. The deceased were buried in a flexed position on either the right or left side, with the head resting on a cobblestone. Some burials were surrounded by pottery sherds and cattle bones. An essential type of burial goods of the Kamarakhevi complex is ceramic vessels, typically numbering two or three per tomb, though occasionally only one or as many as four were found. A few graves lacked pottery altogether. The ceramic assemblage primarily reflects domestic usage. Jewellery was commonly included, with beads made from semi-precious stones and glass found especially in higher-status burials. Iron was the primary material for weapons, and some of the bronze bracelets display parallels with finds from sites such as Beshtasheni, Manglisi, Didube, and Tskneti. The burial practices at Kamarakhevi exhibit numerous parallels with those observed at other sites across the Caucasus, particularly in Eastern Transcaucasia. Additionally, the rich archaeological material from the Kazbegi cemetery—including bracelets, rings, and beads—shares stylistic and typological features with the Akhalgori hoard, providing important chronological indicators for the dating of the Kamarakhevi cemetery. The earliest phase of the cemetery (Group 1; 6th -5th centuries BCE) includes burial complexes that contain ceramic vessels, bracelets decorated with snake heads, buckles with prominent heads, seals with images typical of the Achaemenid period, bent iron knives, and various types of beads, including sard beads. The second group (5th -4th centuries BCE) includes the greatest number of burials, and encompasses the full range of Kamarakhevi pottery, fittings with finely crafted heads, arc-shaped bracelets made of both bronze and iron, iron-tongued bow-shaped strap ends, curved iron knives, and iron spearheads. The third group, attributed to the 4 th -3rd centuries BCE, features cist graves with individuals interred on their right or left side, in a flexed position. Burials from this phase show a significant reduction in bronze jewellery, the absence of sard beads, and a shift to iron for military and agricultural tools. The ceramic products from this period represent a relatively late stage of craftsmanship. Tomb 40. The dimensions of the tomb are: a) 17. X; LI. 0. 1,40X0.86; b) 1,10X0,70 m. The deceased is on the right. A knife and iron fragments have been found. 1082. Tomb 40. Exterior view. 1083. Interior view. მასალები ქართლის სამეფოს ისტორიისთვის ძვ.წ პირველი ათასწლეულის მეორე ნახევარში (კამარახევის სამაროვანი). თბილისი 1960 Materials for the history of the Kingdom of Kartli in the second half of the first millennium BC (Kamarakhevi burial ground) dissertation/ R. Ramishvili; Academy of Sciences of the Georgian SSR; Iv. Javakhishvili Institute of History, 1960. Ramishvili, R. Kamarakhevi Cemetery. in Materials for the Archeology of Georgia and the Caucasus, Vol. 2 (Tbilisi: Development of the Academy of Sciences of the Georgian SSR,1959). Skourtanioti, Eirini, Xiaowen Jia, Nino Tavartkiladze, Liana Bitadze, Ramaz Shengelia, Nikoloz Tushabramishvili, Gunnar U. Neumann, Raffaela Angelina Bianco, Angela Mötsch, Kay Prüfer, Thiseas C. Lamnidis, Luca Traverso, Claudia Sagona, Luka Papac, Wolfgang Haak, David Reich, Sturla Ellingvåg, Philipp W. Stockhammer, Johannes Krause, Harald Ringbauer (2025), The Genetic History of the South Caucasus from the Bronze to the Early Middle Ages: 5000 years of genetic continuity despite high mobility, bioRxiv 2024.06.11.597880; doi: https://doi.org/10.1101/2024.06.11.597880 Jgharkava 1983 - Jgharkava T. Kamarakhevi Cemetery. Catalogue. - A. Apakidze (ed.), Mtskheta VI, 139-190. Tbilisi (in Georgian). ჯღარკავა. კამარახევის სამაროვანი. მცხეთა VI. 73 KIPCHAKOVO, RUSSIA. 200 BCE – 100 CE. BELAYA PYANY BOR. I27018, BURIAL 19, MALE INDIVIDUAL MtDNA H2 a1e1 III. The Middle Kama and Cis-Urals (including the Belaya basin) III.A. The Early Iron Age Pyany Bor culture in the Southern Cis-Urals (Belaya_Pyany-Bor) Genetic results. In this group, we analyzed ten individuals from the sites of Starokirgizovo (n=5) and Kipchakovo (n=5). All 10 individuals had sufficient coverage for downstream population genetic modelling (>100,000 SNPs), and 8 were suitable for IBD-sharing analysis (>600,000 SNPs). No close relatedness was detected among these samples, and the female to-male ratio was 1:9. The samples displayed a genetic composition similar to that of the Low Kama region Iron Age groups. The supervised ADMIXTURE analysis revealed elevated levels of Samara_EBA_Yamnaya and Yakutia_LNBA ancestries in this group. The Belaya_PyanyBor samples are predominantly clustered within the Eurasian Steppe Iron Age (IA) cluster. Similar to the Low Kama Pyany Bor groups, these genetic results suggest the presence of a previously undescribed Iron Age population in the region. III.A.1. Kipchakovo burial site The Kipchakovo burial site is located 1 km east of the village of Minnyarovo (Aktanyshsky district, Republic of Tatarstan) and 2.5 km southwest of the village of Kipchakovo (Ilishovsky district, Republic of Bashkortostan), on the main terrace of the right bank of the Syun River, 70-80 m southwest of the outer rampart of the Kipchakovo settlement. Over the course of four field seasons (1990–1991, 1994, and 2001), F.M. Tagirov excavated 4 kurgans (numbered 15, 41, 51, 52) and two adjacent excavations, one near kurgan 41 (Excavation 1) the other near kurgan 15 (Excavation 2). A total of 64 burials were excavated during this time. Later, the site was excavated by Sergei Zubov. We sequenced five individuals from the site, from the graves burial 12 (individual ID I27117, Male), burial 15 (individual ID I27017, Male), burial 19 (individual ID I27018, Male), burial 6 (individual ID I27020, Male), burial 61 (individual ID I27019, Male). Gyuris, Balázs et al. (2025). Long shared haplotypes identify the southern Urals as a primary source for the 10th-century Hungarians. Cell, Volume 188, Issue 21, 6064 - 6078.e11. DOI: 10.1016/j.cell.2025.09.002 Zubov, S. E., Sattarov, R. R., & Tagirov, F. M. (2021). KIPCHAKOVSKY I KURGAN-GROUND GROUND. Archeology of the Eurasian Steppes , (2), 261– 300. https://doi.org/10.24852/2587-6112.2021.2.261.300 Plans of excavations at Kipchakovo I barrow and subsoil burial ground carried out by F.M. Tagirov in 1990, 1991, 1994, 2001 88 CELJA, CELEIAPARK, SLOVENIA. 4th – 5th CENTURY CE. LATE ROMAN. CE-CP_31/ CPK020. MALE MtDNA H2a1e1a Celeia region case study Municipium Claudium Celeia (Celje) was a Roman town in the province of Noricum Mediterraneum, situated on the itinerary road Poetovio-Aquileia between Pannonia and Italy. It flourished between the 1st-4th century CE and was gradually abandoned in the second half of the 5th c. CE. Mariborska cesta III (183 graves) and Celeiapark (36 graves) sites are parts of the northern and eastern necropoleis of Celeia respectively. The sites are not far from the early Christian church and baptistery, Mariborska cesta III is dated to the 4th c. and Celeiapark to the late 4th-5th c. CE. Breg (20 graves) is part of the southern necropolis of Celeia, located on the southern (right) bank of the Savinja river, dated to the second half of the 4th c. CE. (Table S1 and S3) After the town was abandoned the wealthier inhabitants moved either to safer regions in the Adriatic coast or Italy. A part of the late antique population built settlements on hilltops in the hinterground of the Roman towns. Rifnik near Šentjur is one such hilltop site, consisting of a defence wall, several masonry dwelling houses, water cistern and two churches, and a cemetery with 109 graves outside the defence wall. It is a potential location for the translocated people from Celeia, positioned only about 10 km away. Mariborska cesta III, Celeiapark, Breg In the northern and eastern necropoleis of Celeia (archaeological sites Mariborska cesta III and Celeiapark) in 88 graves 100 individuals were buried. Most of the graves were singular, while two to four individuals were buried together in the rest of the graves. Additionally, at Celeiapark a mass grave with 21 skulls was discovered. Thus, altogether, 121 individuals were documented. One third of the individuals were non-adults, while two thirds were adults with 60:40 ratio males vs. females. In the mass graves, one subadult male and four females were present, twelve were adult males and five undetermined. Pathological changes observed indicate various dental and joint diseases, possible lack of nutrients and rarely healed fractures, neoplasms or inflammations. Out of the 60 discovered individuals buried in 57 graves, 24 individuals (40%) were sampled from Mariborska cesta III. All the individuals sampled were adults, 10 were anthropologically females (42%) and 14 were males (58%). Out of 40 individuals buried in 31 graves at Celeiapark, 22 were sampled. One non-adult individual of undetermined sex and 21 adults, 10 females (48%), 8 males (38%) and three individuals (14%) of undetermined sex were sampled. At Breg, 13 graves (15 skeletons) had been discovered in 1955 and 2 more in 2020, but the bones are not preserved. 20 graves were excavated in 2010 and out of those 21 individuals could be sampled from this site. All the graves were singular, with the exception of graves 1 (two adult females) and 6 (40 weeks in utero child (not sampled) next to an adult female). 15 individuals (71%) were adults and five were non-adults (24%), predominantly children younger than 10 years of age, and one juvenile individual (5%). Anthropologically, eight (53.3%) adult individuals were assessed as female and five (33.3%) as males, two (13.3%) were undetermined. Ratej, R. (2018). Poznoantično grobišče »Celeiapark«. – Magistrsko delo. Oddelek za arheologijo, Filozofska fakulteta, Univerza v Ljubljani, Ljubljana Vyas et al. (2025). The shifting dynamics of ancestry and culture at a post-Roman crossroads. bioRxiv 2025.12.12.693924; doi: https://doi.org/10.64898/2025.12.12.693924 Figure: Grave 31 (Foto: Arhej - Arhiv Celeiapark, Ratej, R. 2018) 89 – 90 ELS MASOS, TARRAGONA, CAMBRILS, SPAIN. LATE ROMAN. 200 BCE - 500 CE. I29526, FEMALE MtDNA H2a1e1a - I29524, FEMALE MtDNA H2a1e1a. Tarragona, Cambrils, Els Masos The area where the site is located has traditionally been used for dry farming. The remains of a residence were found, which likely served as the center of an agricultural estate. It had a luxurious residential section with a peristyle, private baths, and decorations including paintings, mosaics, and marble stonework. The settlement was founded in the mid-1st century CE and lasted until the last quarter of the 3rd or first quarter of the 4th century, at which point the villa was abandoned. In October 1999, under the direction of Ester Ramon from Arqueologia i Patrimoni, an archaeological intervention took place during the construction of a water pipeline to supply the Ponent residential areas (Camarasa, 2002; Ramon, 1999). During this intervention, several architectural structures were identified, highlighting the significance of the settlement. In 2005 and 2006, during the construction of the A-34 highway, another archaeological intervention was conducted, directed by Pèir Còts from SICOM INFORMÀTICA SL, involving the excavation of an area of about 20,000 square meters (Cóts, 2005b, 2005a). It's located on the coastal plain, about 20 kilometres southwest of Tarragona, on the left bank of the Riu de canyes stream, 2 kilometers from the current coastline, within the municipality of Cambrils (Baix Camp). We sampled three individuals, deposited in the Museum of Cambrils, who correspond to a father-mother-daughter trio. Unfortunately, the excavation report has not been made available, which is why the details of the grave(s) are unknown. Graves: • Genetic Identifier: I29524. Grave Identifier: 3012. Grave Type: untraceable. Skeletal information: untraceable. Grave goods: untraceable. Dating: Archeologically dated to 200 BCE - 500 CE. Ancestry summary: Iberia_IA__+__WestAnatolia_Roman_Byzantine__+__North_African_Punic. • Genetic Identifier: I29525. Grave Identifier: 3076. Grave Type: untraceable. Skeletal information: untraceable. Grave goods: untraceable. Dating: Archeologically dated to 200 BCE-500 CE. Ancestry summary: Iberia_IA__+__WestAnatolia_Roman_Byzantine__+__North_African_Punic. • Genetic Identifier: I29526. Grave Identifier: 3256. Grave Type: untraceable. Skeletal information: untraceable. Grave goods: untraceable. Dating: Archeologically dated to 200 BCE-500 CE. Ancestry summary: Iberia_IA__+__WestAnatolia_Roman_Byzantine__+__North_African_Punic. Carrión, P. et al. (2024). Disparate demographic impacts of the Roman Colonization and the Migration Period in the Iberian Peninsula. bioRxiv 2024.09.23.614606; doi: https://doi.org/10.1101/2024.09.23.614606 Figure: View of the roman villa de Els Masos. Pilae of the its bath. La Villa Romana de la Llosa, Guia Arqueologica Visual 96 BALINTORE, SCOTLAND. PICT CULTURE. 419-538 cal CE. INDIVIDUAL BAL003/ INVMG 1982.083. FEMALE MtDNA H2a1e Balintore Balintore today is a village in northern Scotland located in the Highland council region. In 1982, developments near the village (Fearn parish) led to the discovery of two extended inhumations whilst excavating a trench for a new sewer pipe (NRHE 15248) (1). The bodies (accession numbers INVMG1982.082 [listed erroneously as INVMG1985.082 in Sheridan et al. (2)] and INVMG1982.083) were both laid out in long cists in a southwestnortheast position with the head to the southwest. No grave goods were associated with the bodies. Radiocarbon dates suggest the burials date to the 5th-7th centuries AD. Other reports of extended inhumations such as NRHE 15259 suggest a larger cemetery, although it should be noted that some possible Pictish burials from Balintore have been determined via radiocarbon dating to be Neolithic in date (3). Balintore is located close to Shandwick where an intricately carved Pictish cross-slab of 8th-9th century date suggests some form of high status ecclesiastical or secular focus in the later first millennium AD. “BALINTORE (Fearn p) C Millar, D Ross, G Harden Extended Inhumations NH 866 758 In May 1982 two extended inhumations were discovered whilst excavating a trench for a new sewer pipe. They appeared to have been laid out SW-NE with the head at the SW, and had been covered by a number of flat slabs laid across each body. There were no associated finds. The skeletal material is in Inverness Museum, INVMG 982.82 & 83. Sponsor: Inverness District Council” Millar C, Ross D, Harden G. Balintore (Fearn p). In: Discovery and excavation in Scotland. Proudfoot, E.V.W. The Council for British Archaeology, Scotland; 1985. p. 23. Morez A, Britton K, Noble G, Günther T, Götherström A, et al. (2023) Imputed genomes and haplotype-based analyses of the Picts of early medieval Scotland reveal fine-scale relatedness between Iron Age, early medieval and the modern people of the UK. PLOS Genetics 19(4): e1010360. https ://doi.org/10.1371/journal.pgen.1010360 Figure: View of Balintore. Pictish stone collection, Inverness Museum and Art Gallery © Ewen Weatherspoon 97 BRÜCKEN, BRÜCKEN-HACKPFÜFFEL, SAXONY-ANHALT, GERMANY. THURINGIAN KINGDOM. FEMALE MtDNA H2a1e1a “The early medieval burial ground at Brücken, located in the Mansfelder Land district, was thoroughly investigated and completely recorded in 2020 in preparation for a construction project. The site is situated east of district road 2298, approximately 1.3 kilometres south of the village of Brücken, covering an excavation area of 6,525 m². Comprehensive documentation of the excavation is available. A total of 76 graves containing 79 individuals of various ages and sexes were uncovered. Additionally, a sacrificial pit was found containing 5 horses, 4 cattle, and 2 dogs, all of which were killed relatively young, aged 2-5 years, and were predominantly male (analysis conducted by Carola Oelschlägel). The skeletal remains were subjected to anthropological, and radiometric analyses. However, about a third of the human bones analyzed for radiocarbon dating at the Curt-Engelhorn-Zentrum Archäometrie in Mannheim required a second sampling due to low collagen content. Samples were also taken from the human remains for genetic and isotope analyses. Consequently, 12 individuals could not be dated or sampled due to their poor preservation and lack of bone tissue. Initial assessments of the 14C values and characteristic grave goods suggest a Germanic-Thuringian utilization phase of the site between approximately 460 and 540 CE. The grave inventories indicate a clear social hierarchy among the interred, ranging from valuable weapons and jewellery to inferior working tools and, in some cases, no grave goods at all. These findings, combined with the period of burial, suggest that the site served as a burial ground for the inhabitants of a manorial farmstead. The grave goods, including armour, household items, amulets, and clothing accessories, feature local Thuringian material culture as well as objects from distant regions. For instance, Frankish tableware and Alemannic jewellery suggest contacts with the Rhineland and southwestern Germany. The Brücken cemetery's size is typical for burial grounds in the early medieval Thuringian kingdom (c. 455-531). A comparison with other published Thuringian cemeteries—though many have not been fully documented—shows that more than one hundred burials are rarely found79. About 60 cemeteries and burial sites from this period are known in Saxony-Anhalt, but only the Brücken necropolis has been fully identified and extensively documented. This dataset provides a unique opportunity for demographic calculations based on reliable figures for a localized settlement. Moreover, the results of the strontium isotope and genetic analyses can verify the extent to which traditional ethnographic interpretations (typology, stylistics, find mapping) correlate with the origin and ancestry of the individuals. The associated farmstead has not yet been discovered. However, based on settlement studies 80,81 and the terrain, it was likely located within a radius of approximately 500-1,000 meters. The location was strategically chosen, lying between the Harz Mountains and the Kyffhäuser Mountains within the Golden Aue, an area known for its fertile soils, favourable climate, moderate terrain, and good water supply82. The site is on a ridge protected from flooding, within a corridor about three kilometres wide between the Pfüffeler Bach to the south and the small river Helme to the north. Records from Carolingian and Ottonian sources indicate that several long-distance roads converged in this area, meeting near the modern village of Brücken to cross the once marshy Helme and connect with a military road running northeast. This strategic location likely played a crucial role in the settlement’s establishment, providing access to rare goods from abroad. The anthropological analysis of the skeletal remains is limited by their predominantly poor to moderate preservation, affecting age determination, sex assessment, and the identification of pathological conditions. Overall, 29% of the individuals were subadults, including the infans and juvenile age classes. Younger adults (20-40 years) constituted 35%, while older adults (40-60 years) made up 36%. Morphological sexing indicated that 44% of the adults were certainly or probably female, whereas only 22% were determined to be male or probably male. However, genetic sex determination revealed discrepancies regarding the individuals identified as probably female. Some poorly preserved skeletal remains, initially classified as potentially female, were genetically identified as male. This discrepancy is partly due to the poor preservation of the pelvis and the general gracility of the long bones and skulls. Consequently, the gender ratio shifts to a more balanced distribution: 32% female and 29% male. Pathological analysis revealed a high frequency of dental diseases in 51% of individuals, including calculus, periodontal disease, caries, and apical processes. Additionally, degenerative diseases, particularly of the spine, were detected in 29% of older adults. Healed traumas were present in 5% of the postcranial skeleton and 2.5% of the skulls. A notable finding is an artificially deformed skull from feature 46, identified in a robbed grave of a 45- to 55-year-old man, whose sex was also genetically confirmed. This individual is currently the only man with an artificially deformed skull found in Eastern Germany. The tomb is made coffin, containing a high-status 30-40 years old woman carrying MtDNA H2a1e1a, laid in supine position. She was buried with 2 Bow brooches, 1 bird brooch, a gold bracteate, bear tooth pendant, crystal whorl, 17 glass beads, 1 amber bead and eggs. “ Gretzinger, J., Biermann, F., Mager, H. et al. Ancient DNA connects large-scale migration with the spread of Slavs. Nature (2025). https://doi.org/10.1038/s41586-025-09437-6 Figure: Bird brooch, landesmuseum-vorgeschichte, 2021. 101 APPIANO SAN PAOLO CASTELVECCHIO/EPPAN ST. PAUL / ALTENBURG HILL, ALTO ADIGE, ITALY. 650-780 CAL. AD, LANGOBARD FRANKISH. AP AL1, MOL ID 2273, FEMALE INDIVIDUAL MTDNA H2a1e In the municipal area of Appiano, numerous late Roman and early medieval finds have been found in the sites of Lambrecht and Predonico/Perdonig. Furthermore, excavations in recent years, especially in the hamlet of San Paolo, have also yielded significant findings. In 2005, during an inspection of a building project, a large Roman villa with bathing system, featuring mosaic floors (one polychrome and two with black and white tesserae), and wall paintings has been discovered. This is a structure of uncommon wealth considering the local archaeological situation. Archaeological evidence from the early Middle Ages came to light in 2009 in the center of Appiano and on the Altenburg hill. In the center of Appiano, in the basement of a house, approximately 50 meters from the church of San Michele, traces of human habitation were uncovered during renovation work. The excavations, conducted on a very limited area, did not reveal any walls, but did reveal layers with artifacts from the Roman era and the early Middle Ages, including a fibula in the shape of an animal (horse, griffin?) attributable to the 6th-7th century. On the Altenburg hill, close to the center of the hamlet of San Paolo, recoveries of mainly prehistoric ceramic material have been made in recent years. The discovery of a human skeleton triggered a campaign of excavation. On the southern slope of the hill, approximately halfway up, 50 cm deep under the turf, two burials were discovered inserted into the rubble of a probable prehistoric rampart. They did not contain any grave goods. A radiocarbon dating was carried out at the University of Lecce, which indicated an age between 650-780 AD (83.8%). This provides the first clear evidence to support the settlement of this hill in the early Middle Ages. The female individual carrying the MtDNA H2a1e (AP-AL1) has been found in the Altenburg hill (altitude 395 m, sector 4,4C,3A). She was a female adult aged of 30-35 years old. She was dated from the 7th-8th AD which coincides with the Phase 2 of the peopling of the area. This Phase 2 started with the Frankish military expedition mentioned by Paul the Deacon in his Historia Langobardorum, during which many Late Roman forts or places have been destroyed (590 AD). The MtDNA H2a1e is already expanding from Central Europe towards Western Europe during Bronze Age and this lineage could be already installed in the Adige area before this phase. Figure: Map of South Tyrol showing the valleys and the archaeological sites investigated (2) Appiano, San Paolo Castelvecchio (Eppan, St. Paul Altenburg). Map made with the support of Dr. Kathrin Renner (tool: ESRI ArchGIS). Coia, V. & Paladin, A. & Zingale, S. & Croze, M. & Cavada, E. & Lösch, S. & Maixner, F. & Zink, A. (2022). Mitogenomic diversity and stable isotopes provide insights into the maternal genetic history, mobility patterns, and diet of early medieval individuals from the Eastern Italian Alps. Archaeological and Anthropological Sciences. 14. 10.1007/s12520-022-01679-z. LUNZ R., 1990 - Ur- und Frühgeschichte des Eppaner Raumes, Katalog zur archäologischen Ausstellung: 7 April - 9 Mai 1990. Fabio Giovannini, Studi di resti scheletrici umani d’età tardoantica rinvenuti in Alto Adige: malattie, alimentazione, aspetti demografici ed etnici, in: Lorenzo Dal Ri / Stefano di Stefano (a cura di), Archeologia Romana in Alto Adige: Studi e contributi (Beni culturali in Alto Adige – Studi e ricerche I), Bolzano–Vienna 2002, 1091–1105, i. p. 1099–1102. Paladin A, Zink A (2015). Studio antropologico e paleopatologico dei resti scheletrici umani altomedievali del sito di Appiano-Eppan località Langhütten-Gand (Bolzano). In G Kaufmann (Editor). Archäologie des Überetsch, Archeologia dell’Oltradige. Universitätsverlag Wagner, Innsbruck, pp. 391-403. Marzoli C, Bombonato G, Rizzi G (2009) Nuovi dati archeologici sull’insediamento tardo antico-altomedievale della Valle dell’Adige tra la Merano basin e Salorno. Atti Acc. Rov. Agiati, 259 ser. VIII, vol IX, A, fasc. II, 1: 145–183 Marzoli c., 2009 - Appiano/San Paolo, Castelvecchio (G.p. 2484, K.G. Eppan), in «Tutela dei beni culturali in Alto Adige 2008», Lana, pp. 174-175. 106 GANDIA, SANXO LLOP (VALENCIA, SPAIN). 668-874 CE. LATE VISIGOTHIC. GOG35 (Pit N°50). FEMALE MtDNA H2a1e1a The remains of an adult (GOG34) and an infant (GOG35) from the late Visigothic period in Spain (668–874 cal. CE) (Figure 1A, Table 1) were found together in a round pit excavated in a site near Gandía (Figure 1A, D). Genetic analysis identified the adult (GOG34) as male and the infant (GOG35) as female (Table S15). They carried MtDNA haplogroups HV+16311 and H2a1e1a, respectively. The adult carried a R1b1a1b1a1a-L11 (likely unresolved) Y-chromosome (chrY) haplogroup. READ (Figure S3) detected a first-degree kinship between these two individuals. Combined with their ages at death, simultaneous burial, sex identification and non-matching MtDNA haplogroups, we inferred the kinship status to be father–daughter “The stratigraphic record indicates that both bodies were buried in a narrow time frame, since both show the same topographic location, both are deposited on the same base of the silo and both are covered by the same stratum, without there being any type of sediment between the bones of both subjects. To determine the relative chronology of the two depositions, we must pay attention to the fact that the individuals are in contact by means of particularly weak elements. Both the axial and appendicular skeletons of both individuals were well articulated, preserving both persistent and weak anatomical connections. Therefore, the corpses had all the anatomical integrity at the moment in which they came into contact and the disposition of one has not led to the alteration of the other. It could therefore be a simultaneous burial or two inhumations carried out within a short period of time. It is difficult to be more precise. However, studying the reciprocal positions the simultaneity of the deposition can be deduced. The distribution of the two bodies in the grave also supports the assumption that both bodies were buried at the same time. Both deceased seem placed respecting the space that the body volume occupies, one from the other. In this sense, the position of the lower extremities of the adult individual compressed against the wall of the silo, with the articulation of the knees fully flexed, seems to obey the need to leave enough space for the deposition of the infantile individual The paleopathological analysis did not reveal a violent traumatic injury or an infectious disease that could have caused the death of both in a very short space of time. The adult individual presents several traumatic injuries such as trauma marks in the lumbar vertebrae, signs of fracture of the left clavicle and posttraumatic myositis ossificans in the distal third of the right tibia, on the insertion surface of the interosseous membrane with the fibula, at the level of the opening for the perforating branch of the peroneal artery. However, all these injuries were consolidated long before death and are undoubtedly caused by hard and continuous physical activity. Regarding the pathologies detected in the subadult individual, the incisor teeth displayed banded depressions Fig. 3. Plan of the UE50 burial and representation of the original position in which the bodies were buried. P. Mas and Ll. Alapont. Fig. 4. Post-traumatic myositis ossificans in the distal third of the right tibia of the male adult individual. in the dental enamel, a physical indicator of hypoplasia. This occurs during dental development and remains throughout life as proof of having suffered disease or malnutrition during childhood. An indication of possible growth and biological development problems. In addition, the child had multiple holes in the eye sockets. These holes are evidence of porotic orbital sieve. This pathology is associated with anaemia and other diseases related to a deficient assimilation of vitamins and nutrients. Anaemia is the result of a lack of iron that can be genetically inherited or a consequence of a lack of iron in commonly consumed foods. The diseases observed in the infantile subject can be related to some type of epidemic or endemic infectious disease. Oteo-García G, Alapont-Martín L, Pascual Beneyto J, Foody MGB, Yau B, Pala M, et al. Late Roman tombs at Sanxo Llop (Gandía, Valencia): Exogamy and kinship in a particular funerary structure. In: Death and the Societies of Late Antiquity: New methods, new questions? (Archéologies méditerranéennes). Aix-en-Provence: Presses universitaires de Provence; 2023. p. 119–27. Oteo-Garcia G., Marina Silva, M. George B. Foody, Bobby Yau, Alessandro Fichera, Llorenc Alapont, Pierre Justeau, Simao Rodrigues, Rita Monteiro, Francesca Gandini, Marisa Rovira, Albert Ribera i Lacomba, Josep Pascual Beneyto, Valeria Mattiangeli, Daniel G. Bradley, Ceiridwen J. Edwards, Maria Pala, Martin B. Richards (2024). Medieval genomes from eastern Iberia illuminate the role of Morisco mass deportations in dismantling a longstanding genetic bridge with North Africa bioRxiv 2024.10.09.617385; doi: https://doi.org/10.1101/2024.10.09.617385 Figure 34: Plan of the UE50 burial and representation of the original position in which the bodies were buried. P. Mas and Ll. Alapont. 123 GROENMARKT-2, SINT-TRUIDEN, BELGIUM. 10TH - 11TH C. AD. SK2937 / ST2937/ LAYER 11, FEMALE INDIVIDUAL MtDNA H2a1e1a SK2922, SK2923, SK2936, SK2937, SK2938, SK2940 (WP 2, zone 2, VL 14): southwest/northeast oriented grave containing the remains of three adults, two children, and one juvenile. Based on their physical characteristics, the adults all appear to be male; the children (SK2936 and SK2937) are approximately six and five years old, respectively. DNA analysis has identified SK2937 as female. SK2922, located at the very top of the grave, has been dated using radiocarbon dating. The skeleton dates to the 10th century or the first quarter of the 11th century; this grave is therefore certainly earlier than the construction of the Church of Our Lady under Abbot Adélard. Beneker, O., Molinaro, L., Guellil, M. et al. Urbanization and genetic homogenization in the medieval Low Countries revealed through a ten-century paleogenomic study of the city of Sint-Truiden. Genome Biol 26, 127 (2025). https://doi.org/10.1186/s13059-025-03580-z de Winter, N. (2023). ARON rapport 1258 – Eindverslag Sint-Truiden Groenmarkt. Opgraving naar aanleiding van de herinrichting van de Groenmarkt, het Trudoplein, de Diesterstraat, de Plankstraat en de Meinstraat. Tech. Rep. 1258 ARON. De Winter, N. (2024). De Winter N. 2024: vondst van drieduizend graven in het stadscentrum van Sint-Truiden, M&L jg.43/1, p.27-42. M&L. 134 CHERRY HINTON, CAMBRIDGESHIRE (RURAL PARISH ). 940-1170 AD. LATE MEDIEVAL. SKL ID 2256; BURIAL 70; ID 906 , FEMALE INDIVIDUAL MtDNA H2a1e1a Cherry Hinton The settlement of Church End Cherry Hinton (Cherry Hinton) is located around six kilometers southeast of Cambridge. In the late 9th to the mid10th century, a large thegnly (aristocratic) or proto-manorial center was established. The associated timber chapel and graveyard were excavated in 1999 by the Hertfordshire Archaeological Trust (subsequently Archaeological Solutions and now Wardell Armstrong) in advance of development of the site in accordance with the appropriate planning regulations. The burials have been dated by a combination of stratigraphy, typological dating of associated artefacts particularly pottery, art-historical dating of associated sculpture, parallels for the associated church and radiocarbon dating of a sample of the skeletons. The skeletons were initially studied by Lavinia Ferrante di Ruffano and Tony Waldron for the Hertfordshire Archaeological Trust and were reexamined for the After the Plague project by Sarah Inskip. The human skeletal remains are held by the Historic Environment Team of Cambridgeshire County Council ([email protected]), to whom any requests for access should be made. Only part of the cemetery was investigated, including over 670 graves and the remains of c. 980 individuals. Most burials were west-east aligned extended supine inhumations, with the head to the west. The cemetery population was estimated to be c. 1000–2000. The cemetery probably served an entire thriving Late Saxon to Norman rural settlement, broadly representative of a mixed rural peasant tenant hierarchy engaged primarily but not exclusively in agricultural labor. The burials follow a relatively uniform rite, so there are few archaeological indications of the status of particular individuals. Based on radiocarbon dating plus a range of other evidence the cemetery dates to c. 940/990–1120/70. As the population from which those buried at Church End were drawn probably increased over time it is likely that the majority, possibly around two thirds, died after the Norman Conquest. As this settlement was located firmly within the rural hinterland of Cambridge this population provides an important comparator to the town, albeit one that is rather earlier than the main populations that have been studied from there. In total, 48 individuals from Cherry Hinton were targeted for DNA extraction in this study, including 24 females and 24 males. C. Cessford, A. Dickens, The Manor of Hintona: The origins and development of Church End, Cherry Hinton. Proc. Camb. Antiqu. Soc. 94, 51–72 (2005). C. Cessford, A. Slater, Beyond the manor of hintona further thoughts on the development of church end, cherry hinton: The neath farm site. Proc. Camb. Antiqu. Soc. 103, 39–60 (2014). M. Lally, 69 to 115 Church End, Cherry Hinton, Cambridgeshire: Post Excavation Assessment and Updated Project Design (Archaeological Solutions Report 3012, 2008). Hui, R. et al. (2023). Medieval social landscape through the genetic history of Cambridgeshire before and after the Black Death; BioRxiv 6 mars 2023; t doi: https://doi.org/10.1101/2023.03.03.531048 Figure: Map of excavations (C. Cessford, A. Slater 2014) 135 SANTOK, LUSZ VOIDVOSHIP, POLAND. 1000 -1200 AD. PCA 0377, FEMALE MtDNA H2a1e1a Santok The remains of the vast necropolis in Santok (Lubuskie voivodeship) were discovered in 2014 during archaeological research accompanying the construction of sanitary sewage system. The cemetery is located on the Warta River terrace, at the foot of one of the two Santok strongholds referred to as Pomeranian, in contrast to the fortified settlement on the other bank of the river, which was part of the Piast State. The excavations were carried out in a 1.4-2 m wide strip, recording skeletal graves in a section of about 430 m. During the rescue excavation, 197 burials were registered, lying on several levels, but the final anthropological analysis showed that as many as 328 individual remains were excavated from the graves. The cemetery dates back to the turn of the 11th and 12th centuries, up to the 13th century. Tools (whetstones, knives), ornaments (rings, temple rings) and single coins were discovered in the burials. In numerous cases, traces of wooden structures, including coffins, were also recorded. Stolarek, I. et al. (2023), Genetic history of East-Central Europe in the first millennium CE. Genome Biology (2023) 24:173 https://doi.org/10.1186/s13059-023-03013-9 BIBLIOGRAPHY Allentoft, ME et al. (2015). Population genomics of Bronze Age Eurasia. Nature. 2015 Jun 11;522(7555):167-72. Doi: 10.1038/nature14507. PMID: 26062507. Andreeva, Tatiana V. et al. (2025). Genetic history of Rus’.bioRxiv 2025.12.30.695215; doi: https://doi.org/10.64898/2025.12.30.695215 Anthony D. W., Khokhlov A. A., Agapov S. A., Agapov D. S., Schulting R., Olalde, I., Reich D. (2022). The Eneolithic cemetery at Khvalynsk on the Volga River, Praehistorische Zeitschrift 2022; aop. De Gruyter. https://doi.org/10.1515/pz-2022-2023 Antonio, M. L. Et al. (2019). Ancient Rome: A genetic crossroads of Europe and the Mediterranean. Science 08 Nov 2019: Vol. 366, Issue 6466, pp. 708-714 DOI: 10.1126/science.aay6826 Barrie, W., Yang, Y., Irving-Pease, E.K. et al. Elevated genetic risk for multiple sclerosis emerged in steppe pastoralist populations. Nature 625, 321– 328 (2024). https://doi.org/10.1038/s41586-023-06618-z Ben Krause-Kyora et al. (2018). Ancient DNA study reveals HLA susceptibility locus for leprosy in medieval Europeans. Nat Commun 9, 1569 (2018). https ://doi.org/10.1038/s41467-018-03857-x Beneker, O., Molinaro, L., Guellil, M. et al. Urbanization and genetic homogenization in the medieval Low Countries revealed through a ten-century paleogenomic study of the city of Sint-Truiden. Genome Biol 26, 127 (2025). https://doi.org/10.1186/s13059-025-03580-z Brandt, G., et al. (2013), Ancient DNA reveals key stages in the formation of central European mitochondrial genetic diversity. Science (New York, N.Y.), 342(6155), 257–261. https://doi.org/10.1126/science.1241844. Brandstätter, A. et al. (2006). Dissection of mitochondrial superhaplogroup H using coding region SNPs. Electrophoresis 27, 2541-2550, doi: 10.1002:elps.200500772. Brisighelli, F. et al. (2012) Uniparental Markers of Contemporary Italian Population Reveals Details on Its Pre-Roman Heritage. PloS ONE7(12): e50794. Doi:10.1371/journal.pone.0050794 Brotherton, P., Haak, W., Templeton, J. et al. Neolithic mitochondrial haplogroup H genomes and the genetic origins of Europeans. Nat Commun 4, 1764 (2013). https://doi.org/10.1038/ncomms2656 Brunel, S. et al. (2020). Ancient genomes from present-day France unveil 7,000 years of its demographic history. Proceedings of the National Academy of Sciences. 117. 201918034. 10.1073/pnas.1918034117. Burger et al. (2020). Low Prevalence of Lactase Persistence in Bronze Age Europe Indicates Ongoing Strong Selection over the Last 3,000 Years, Current Biology (2020), https://doi.org/10.1016/j.cub.2020.08.03 Cassidy, L.M., Russell, M., Smith, M. et al. Continental influx and pervasive matrilocality in Iron Age Britain. Nature 637, 1136–1142 (2025). https://doi.org/10.1038/s41586-024-08409-6 Carrión, P. et al. (2024). Disparate demographic impacts of the Roman Colonization and the Migration Period in the Iberian Peninsula. bioRxiv 2024.09.23.614606; doi: https://doi.org/10.1101/2024.09.23.614606 Chyleński, M., Makarowicz, P., Juras, A. et al. (2023), Patrilocality and hunter-gatherer-related ancestry of populations in East-Central Europe during the Middle Bronze Age. Nat Commun 14, 4395 (2023). https://doi.org/10.1038/s41467-023-40072-9 Coia, V. & Paladin, A. & Zingale, S. & Croze, M. & Cavada, E. & Lösch, S. & Maixner, F. & Zink, A. (2022). Mitogenomic diversity and stable isotopes provide insights into the maternal genetic history, mobility patterns, and diet of early medieval individuals from the Eastern Italian Alps. Archaeological and Anthropological Sciences. 14. 10.1007/s12520-022-01679-z. Damgaard, P.d.B. et al. (2018). 137 ancient human genomes from across the Eurasian steppes. Nature 557, 369–374 (2018). https://doi.org/10.1038/s41586-018-0094-2 Der Sarkissian, C. (2011). Mitochondrial DNA in ancient human populations of Europe. Thesis (Ph.D.),University of Adelaide, School of Earth and Environmental Sciences. Derenko, M., Malyarchuk, B., Grzybowski, T., Denisova, G., Dambueva, I., Perkova, M., Dorzhu, C., Luzina, F., Lee, H. K., Vanecek, T., Villems, R., & Zakharov, I. (2007). Phylogeographic analysis of mitochondrial DNA in northern Asian populations. American journal of human genetics, 81(5), 1025– 1041. https://doi.org/10.1086/522933 Doan, K., Panagiotopoulou, H., Molak, M., Siekiera, A., Bogdanowicz, W. (2018). Analiza genetyczna szczątków ludzkich, in Cieple. Pp 447-460. Muzeum Archeologiczne w Gdańsku. (2018) Doris Pany-Kucera et al. (2020). Social Relations, Deprivation and Violence at Schleinbach, Lower Austria. Insights from an Interdisciplinary Analysis of the Early Bronze Age Human Remains. Archaeologia Austriaca, Band 104/2020, 13–52 © 2020 by Österreichische Akademie der Wissenschaften, Wien doi: 10.1553/archaeologia104s13 Dulik, Matthew C, (2011). A molecular anthropological study of Altaian histories utilizing population genetics and phylogeography. Dissertations available from ProQuest. AAI3463002. https://repository.upenn.edu/dissertations/AAI3463002 Emery, M. (2018). Ancient Roman mitochondrial genomes and isotopes reveal relationships and geographic origins at the local and panMediterranean scales. Journal of Archaeological Science: Reports, 20(?), 200-209. https://doi.org/10.1016/j.jasrep.2018.04.036 Dániel Gerber et al. ,Ancient genomes reveal Avar-Hungarian transformations in the 9th-10th centuries CE Carpathian Basin.Sci. Adv.10,eadq5864(2024).DOI:10.1126/sciadv.adq5864 Ghalichi, A., Reinhold, S., Rohrlach, A.B. et al. The rise and transformation of Bronze Age pastoralists in the Caucasus. Nature (2024). https://doi.org/10.1038/s41586-024-08113-5 Gnecchi-Ruscone, G.A et al. (2024), Network of large pedigrees reveals social practices of Avar communities. Nature 629, 376–383 (2024). https://doi.org/10.1038/s41586-024-07312-4 Gnecchi-Ruscone, G. A. Et al. (2021), Ancient genomic time transect from the Central Asian Steppe unravels the history of the Scythians. Science Advances 26 Mar 2021: Vol. 7, no. 13, eabe4414 DOI: 10.1126/sciadv.abe4414 Gretzinger, J., Sayer, D., Justeau, P. et al. The Anglo-Saxon migration and the formation of the early English gene pool. Nature (2022). https ://doi.org/10.1038/s41586-022-05247-2 Hernández, C.L., Reales, G., Dugoujon, J. et al. (2014). Human maternal heritage in Andalusia (Spain): its composition reveals high internal complexity and distinctive influences of MtDNA haplogroups U6 and L in the western and eastern side of region. BMC Genet 15, 11 (2014) doi:10.1186/14712156-15-11 Hui, R. et al. (2023). Medieval social landscape through the genetic history of Cambridgeshire before and after the Black Death; BioRxiv 6 mars 2023; t doi: https://doi.org/10.1101/2023.03.03.531048 Järve, M. et al. (2019). Shifts in the Genetic Landscape of the Western Eurasian Steppe Associated with the Beginning and End of the Scythian Dominance. Curr Biol. 2019;29(14):2430-2441.e10. doi:10.1016/j.cub.2019.06.019 Jankova-Ajanovska, R. et al. (2014). Mitochondrial DNA control region analysis of three ethnic groups in the Republic of Macedonia. Forensic Science International Genetics.2014;13:1-2. Doi:10.1016/j.fsigen.2014.06.013 Jeran, N. et al. (2009). Mitochondrial DNA heritage of Cres Islanders—example of Croatian genetic outliers. Collegium antropologicum. 33. 1323-8. Juras et al. (2021). Maternal genetic https://doi.org/10.1002/ajpa.24372 origin of the late and final Neolithic human populations from present-day Poland. Juras, A. et al. (2018). Mitochondrial genomes reveal an east to west cline of steppe ancestry in Corded Ware populations. Scientific Reports. 8. 11603. 10.1038/s41598-018-29914-5. Juras, A. (2012), Ethnogenesis of the Slavs in the light of ancient DNA analyses, thesis. Keyser, C. et al. (2020). Genetic evidence suggests a sense of family, parity and conquest in the Xiongnu Iron Age nomads of Mongolia. Hum Genet (2020). https://doi.org/10.1007/s00439-020-02209-4 Klunk, J. (2018). Genetic causes and consequences of ancient plague pandemics: pathogen evolution and human demographic reverberations, Thesis, Macmaster University, Canada. Knipper, C. et al (2017). Female exogamy and gene pool diversification at the transition from the Final Neolithic to the Early Bronze Age in central Europe.” Proceedings of the National Academy of Sciences 114.38 : 10083-10088. Web. 02 Aug. 2020. Krzewińska, M. et al. (2018). Ancient genomes suggest the eastern Pontic-Caspian steppe as the source of western Iron Age nomads. Sci Adv 4 (10), eaat4457. DOI: 10.1126/sciadv.aat4457 Krzewińska, M. et al. (2015).Mitochondrial DNA variation in the Viking age population of Norway. Philosophical Transactions of The Royal Society B Biological Sciences 370(1660) https://doi.org/10.1098/rstb.2013.0384 Lappalainen, T. et al. (2008). Migration waves to the Baltic Sea region. Ann Hum Genet. 2008 ;72(Pt 3) :337-348. Doi :10.1111/j.14691809.2007.00429.x Lazaridis, I. et al. (2024), The Genetic Origin of the Indo-Europeans, bioRxiv 2024.04.17.589597; doi: https://doi.org/10.1101/2024.04.17.589597 Lazaridis, I. et al. (2022), The genetic history of the Southern Arc: A bridge between West Asia and Europe. Science. 2022 Aug 26;377(6609):eabm4247. Doi: 10.1126/science.abm4247. Epub 2022 Aug 26. PMID: 36007055. Lazaridis, I. et al. (2016). Nadel, D., Rollefson, G. et al. Genomic insights into the origin of farming in the ancient Near East. Nature 536, 419–424. https://doi.org/10.1038/nature19310 Maár, K. et al. (2021). Maternal lineages from 10-11th century commoner cemeteries of the Carpathian Basin. bioRxiv 2021.01.26.428268; doi: https://doi.org/10.1101/2021.01.26.428268 Margaryan, A., Lawson, D.J., Sikora, M. et al. Population genomics of the Viking world. Nature 585, 390–396 (2020). https://doi.org/10.1038/s41586020-2688-8 Maróti, Z. et al. (2022). Whole genome analysis sheds light on the genetic origin of Huns, Avars and conquering Hungarians. BioRxiv, https://doi.org/10.1101/2022.01.19.476915 : Matheson, C. D. et al. (2009) Molecular Exploration of the First-Century Tomb of the Shroud in Akeldama, Jerusalem. PloS ONE 4(12): e8319. https://doi.org/10.1371/journal.pone.0008319 Mathieson, I. et al. (2018), The genomic history of southeastern Europe. Nature 555, 197–203. https://doi.org/10.1038/nature25778 Mathieson, I. et al. (2015), Genome-wide patterns of selection in 230 ancient Eurasians. Nature. 528. 499-503. 10.1038/nature16152. McColl, H., Guus Kroonen, Thomaz Pinotti, John Koch, Johan Ling, Jean-Paul Demoule, Kristian Kristiansen, Martin Sikora, Eske Willerslev (2025). Tracing the Spread of Celtic Languages using Ancient Genomics. bioRxiv 2025.02.28.640770; doi: https://doi.org/10.1101/2025.02.28.640770 McColl, H. et al. (2024), Steppe Ancestry in western Eurasia and the spread of the Germanic Languages, bioRxiv 2024.03.13.584607; doi: https://doi.org/10.1101/2024.03.13.584607 Morez A, Britton K, Noble G, Günther T, Götherström A, et al. (2023) Imputed genomes and haplotype-based analyses of the Picts of early medieval Scotland reveal fine-scale relatedness between Iron Age, early medieval and the modern people of the UK. PLOS Genetics 19(4): e1010360. https ://doi.org/10.1371/journal.pgen.1010360 Nikitin, A.G., Lazaridis, I., Patterson, N. et al. A genomic history of the North Pontic Region from the Neolithic to the Bronze Age. Nature (2025). https://doi.org/10.1038/s41586-024-08372-2 Mittnik, A. et al. (2019). Kinship-based social inequality in Bronze Age Europe. Science. 366. Eaax6219. 10.1126/science.aax6219. Mustafin, K. K. Alborova, I. E. Semenov, A. S. Vishnevsky V. I. (2018). Haplogroup analysus for a medieval Russian burial of 16 th- 17th centuries in Radonezh (Moscow area). УДК 975.174.2 ; ББК 63.2 ; DOI https://doi.org/10.21638/spbu19.2018.209 Narasimhan, V. M. et al. (2019). The formation of human populations in South and Central Asia, Science 365, eaat7487 DOI: 10.1126/science.aat7487 Nazia (2014). Genetic analyses of the major tribes of Abbottabad and Mansehra districts through dental morphology and DNA analyses. Thesis, Department of Genetics, Hazara University Mansehra, 2014. Neparáczki, E. et al. (2017). Genetic structure of the early Hungarian conquerors inferred from MtDNA haplotypes and Y-chromosome haplogroups in a small cemetery. Mol Genet Genomics 292, 201–214 (2017). https://doi.org/10.1007/s00438-016-1267-z Nikitin, Alexey G., Iosif Lazaridis, Nick Patterson, Svitlana Ivanova, Mykhailo Videiko, Valentin Dergachev, Nadiia Kotova, Malcolm Lillie, Inna Potekhina, Marta Krenz-Niedbała, Sylwia Łukasik, Serhij Makhortykh, Virginie Renson, Henry Shephard, Gennadie Sirbu, Sofiia Svyryd, Taras Tkachuk, Piotr Włodarczak, Kim Callan, Elizabeth Curtis, Eadaoin Harney, Lora Iliev, Aisling Kearns, Ann Marie Lawson, Megan Michel, Matthew Mah, Adam Micco, Jonas Oppenheimer, Lijun Qiu, J. Noah Workman, Fatma Zalzala, Swapan Mallick, Nadin Rohland, David Reich (2024), A genomic history of the North Pontic Region from the Neolithic to the Bronze Age, bioRxiv 2024.04.17.589600; doi: https://doi.org/10.1101/2024.04.17.589600 Olalde, I. et al. (2018). The Beaker phenomenon and the genomic transformation of northwest Europe. Nature. 555. 543-543. 10.1038/nature26164. Oteo-García G, Alapont-Martín L, Pascual Beneyto J, Foody MGB, Yau B, Pala M, et al. (2023). Late Roman tombs at Sanxo Llop (Gandía, Valencia): Exogamy and kinship in a particular funerary structure. In: Death and the Societies of Late Antiquity: New methods, new questions? (Archéologies méditerranéennes). Aix-en-Provence: Presses universitaires de Provence; 2023. p. 119–27. Gonzalo Oteo-Garcia, Marina Silva, M. George B. Foody, Bobby Yau, Alessandro Fichera, Llorenc Alapont, Pierre Justeau, Simao Rodrigues, Rita Monteiro, Francesca Gandini, Marisa Rovira, Albert Ribera i Lacomba, Josep Pascual Beneyto, Valeria Mattiangeli, Daniel G. Bradley, Ceiridwen J. Edwards, Maria Pala, Martin B. Richards (2024). Medieval genomes from eastern Iberia illuminate the role of Morisco mass deportations in dismantling a long-standing genetic bridge with North Africa bioRxiv 2024.10.09.617385; doi: https://doi.org/10.1101/2024.10.09.617385 Översti, S. et al. (2019). Human mitochondrial DNA lineages in Iron-Age Fennoscandia suggest incipient admixture and eastern introduction of farming-related maternal ancestry. Scientific Reports. 9, 14 p., 16883.https ://doi.org/10.1038/s41598-019-51045-8 Översti, S. et al. (2017). Identification and analysis of MtDNA genomes attributed to Finns reveal long-stagnant demographic trends obscured in the total diversity. Sci Rep 7, 6193. https://doi.org/10.1038/s41598-017-05673-7 Palanichamy, M. G. et al (2015). West Eurasian MtDNA lineages in India: an insight into the spread of the Dravidian language and the origins of the caste system, Hum Genet DOI 10.1007/s00439-015-1547-4 Papac, L. et al. (2021). Dynamic changes in genomic and social structures in third millennium BCE central Europe. Science Advances, 25 Aug 2021: Vol. 7, no. 35, eabi6941 DOI: 10.1126/sciadv.abi6941 Cassidy, L.M., Russell, M., Smith, M. et https://doi.org/10.1038/s41586-024-08409-6 al. Continental influx and pervasive matrilocality in Iron Age Britain. Nature (2025). Patterson, N., Isakov, M., Booth, T. et al. Large-scale migration into Britain during the Middle to Late Bronze Age. Nature 601, 588–594 (2022). https://doi.org/10.1038/s41586-021-04287-4 Penske S, Küßner M, Rohrlach AB, Knipper C, Nováček J, Childebayeva A, Krause J, Haak W. (2024), Kinship practices at the early bronze age site of Leubingen in Central Germany. Sci Rep. 2024 Feb 16;14(1):3871. doi: 10.1038/s41598-024-54462-6. PMID: 38365887; PMCID: PMC10873355. Penske, S., Rohrlach, A.B., Childebayeva, A. et al. (2023). Early contact between late farming and pastoralist societies in southeastern Europe. Nature (2023). https://doi.org/10.1038/s41586-023-06334-8 Posth, C. et al. (2021). The origin and legacy of the Etruscans through a 2000-year archeogenomic time transect. Sci. Adv. 7, eabi7673 (2021). DOI: 10.1126/sciadv.abi7673 Ravasini, F., Kabral, H., Solnik, A. et al. The genomic portrait of the Picene culture provides new insights into the Italic Iron Age and the legacy of the Roman Empire in Central Italy. Genome Biol 25, 292 (2024). https://doi.org/10.1186/s13059-024-03430-4 Rodríguez-Varela, R. et al. (2023). The genetic history of Scandinavia from the Roman Iron Age to the present, Cell, Volume 186, Issue 1, 2023, Pages 32-46.e19, ISSN 0092-8674, https://doi.org/10.1016/j.cell.2022.11.024 Rym, K. et al. (2016): On the origin of Iberomaurusian: new databased on ancient mitochondrial DNA and phylogenetic analysis of Afalou and Taforalt populations, Mitochondrial DNA Part A, DOI: 10.1080/24701394.2016.1258406 Saag, L. et al. (2020). Genetic ancestry changes in Stone to Bronze Age transition in the East European plain. 10.1101/2020.07.02.184507. Saag, L. et al. (2019). The Arrival of Siberian Ancestry Connecting the Eastern Baltic to Uralic Speakers further East. Current biology: CB, 29(10), 1701– 1711.e16. https://doi.org/10.1016/j.cub.2019.04.026 Skourtanioti, Eirini, Xiaowen Jia, Nino Tavartkiladze, Liana Bitadze, Ramaz Shengelia, Nikoloz Tushabramishvili, Gunnar U. Neumann, Raffaela Angelina Bianco, Angela Mötsch, Kay Prüfer, Thiseas C. Lamnidis, Luca Traverso, Claudia Sagona, Luka Papac, Wolfgang Haak, David Reich, Sturla Ellingvåg, Philipp W. Stockhammer, Johannes Krause, Harald Ringbauer (2024), The Genetic History of the South Caucasus from the Bronze to the Early Middle Ages: 5000 years of genetic continuity despite high mobility, bioRxiv 2024.06.11.597880; doi: https://doi.org/10.1101/2024.06.11.597880 Seguin-Orlando et al. (2020). Heterogeneous Hunter-Gatherer and Steppe-Related Ancestries in Late Neolithic and Bell Beaker Genomes from Present-Day France, Current Biology (2020), https://doi.org/10.1016/j.cub.2020.12.015 Scheib, C. L. et al. (2023), Local population structure in Cambridgeshire during the Roman occupation, bioRxiv 2023.07.31.551265; doi: https://doi.org/10.1101/2023.07.31.551265 Oszkár Schütz, Zoltán Maróti, Balázs Tihanyi, Attila P. Kiss, Emil Nyerki, Alexandra Gînguță, Petra Kiss, Gergely I.B. Varga, Bence Kovács, Kitti Maár, Bernadett Ny. Kovacsóczy, Nikoletta Lukács, István Major, Antónia Marcsik, Eszter Patyi, Anna Szigeti, Zoltán Tóth, Dorottya Walter, Gábor Wilhelm, Réka Cs. Andrási, Zsolt Bernert, Luca Kis, Liana Oța, György Pálfi, Gábor Pintye, Dániel Pópity, Angela Simalcsik, Andrei Dorian Soficaru, Olga Spekker, Sándor Varga, Endre Neparáczki, Tibor Török, Unveiling the origins and genetic makeup of the “forgotten people”: A study of the Sarmatian-period population in the Carpathian Basin, Cell, (2025), ISSN 0092-8674, https://doi.org/10.1016/j.cell.2025.05.009. Stolarek, I. et al. (2023), Genetic history of East-Central Europe in the first millennium CE. Genome Biology (2023) 24:173 https://doi.org/10.1186/s13059-023-03013-9 Stolarek, I., Handschuh, L., Juras, A. et al. (2019), Goth migration induced changes in the matrilineal genetic structure of the central-east European population. Sci Rep 9, 6737. https://doi.org/10.1038/s41598-019-43183-w Tatiana V. Andreeva et al. (2025). Genetic history of Scythia. Sci. Adv.11,eads8179 (2025). DOI:10.1126/sciadv.ads8179 Unterländer, M. et al. (2017). Ancestry and demography and descendants of Iron Age nomads of the Eurasian Steppe. Nat. Commun. 8, 14615 doi: 10.1038/ncomms14615 Vai, S., Brunelli, A., Modi, A. et al. A genetic perspective on Longobard-Era migrations. Eur J Hum Genet 27, 647–656 (2019). https://doi.org/10.1038/s41431-018-0319-8 Vikas K. et al. (2022). Bronze and Iron Age population movements underlie Xinjiang population history; Science, 376 (6588), • DOI: 10.1126/science.abk1534 Wang, CC. et al. (2018),s The genetic prehistory of the Greater Caucasus. BioRxiv; DOI: 10.1101/322347 Wang, CC. et al. (2019), Ancient human genome-wide data from a 3000-year interval in the Caucasus corresponds with eco-geographic regions. Nat Commun 10, 590 (2019). https://doi.org/10.1038/s41467-018-08220-8 Wang, K., Tobias, B., Pany-Kucera, D. et al. Ancient DNA reveals reproductive barrier despite shared Avar-period culture. Nature (2025). https://doi.org/10.1038/s41586-024-08418-5 Wang, W. et al. (2021), mitogenomes reveal intense admixture with high genetic diversity. Sci. Adv. 7, eabd6690 (2021). Wilde S. (2014). Populationsgenetik kupfer- und bronzezeitlicher Bevölkerungen der osteuropäischen Steppe, Johannes Gutenberg-Universität Mainz. Citation: Rondu, M.-O. (2026). MtDNA H2a1 Encyclopedia, MtDNA H2a1e, 2026. Paris. Legal deposit: Empreinte numérique SHA256 43ceda9d283344574744e435e6b60e40aad2f3a6db832392451469f704e42e78