A man buried in Spain in the Copper Age era

I3269
Portrait reconstruction
Specimen Details
Sample ID:
I3269
Date:
3000 BCE - 2800 BCE
Biological Sex:
Male
mtDNA:
U5b2b3a
Y-DNA:
I2a2a
Cultural Period:
Chalcolithic Spain
Location
Country:
Spain
Locality:
Basque Country. Alava. Las Yurdinas II
Coordinates:
Map Location
Historical Timeline
Description

The Chalcolithic period, also known as the Copper Age, represents a transitional phase between the Neolithic (New Stone Age) and the Bronze Age, characterized by the first use of metal tools and weapons. In Iberian Chalcolithic or Chalcolithic Spain, this era is particularly significant as it marks a period of social, technological, and cultural transformation that laid the groundwork for subsequent developments in the Iberian Peninsula.

Chronology and Geography

The Chalcolithic period in the Iberian Peninsula roughly spans from 3000 to 1800 BCE, although the exact dates may vary regionally due to the diversity of cultural developments across the area. The Iberian Chalcolithic culture was widespread across modern-day Spain and Portugal, with notable concentrations in regions such as Andalusia, Extremadura, and central and eastern parts of the peninsula.

Technological Innovations

The hallmark of the Chalcolithic period is the emergence of metalworking, particularly with copper. While stone tools continued to be predominant for numerous functions, the advent of metalwork introduced new tool types, such as axes, blades, and ornaments, indicating both utilitarian and ceremonial uses. The knowledge of metallurgy, although initially basic, laid important foundations for the subsequent Bronze Age.

Settlements and Architecture

Settlements during the Iberian Chalcolithic period became more complex, with a mix of small hamlets, larger fortified villages, and even the first proto-urban centers. One of the most notable sites is Los Millares in Almería, southern Spain, characterized by its impressive fortifications, multiple concentric walls, and complex, densely packed houses. This site, along with others like Vila Nova de São Pedro in Portugal, showcases advanced organizational and construction skills, reflecting emerging social hierarchies.

Social Structure

The Chalcolithic era is marked by increasing social stratification. This is evidenced by the differentiation in grave goods found in burial sites, suggesting a society with distinct social classes and roles. The appearance of megalithic tombs and the prevalence of grave goods indicate the importance of social status, and possibly the emergence of a priestly or ruling class that wielded both religious and secular power.

Economy and Subsistence

Agriculture continued to evolve with a mixed economy of farming and herding. Crops such as barley, wheat, and legumes were commonly cultivated, while the domestication of animals, including cattle, sheep, and goats, remained integral. The Chalcolithic economy also saw an expansion in trade networks, as evidenced by the distribution of exotic materials like amber and ivory, indicating far-reaching contacts across Europe and the Mediterranean.

Cultural and Ritual Practices

Chalcolithic Spain is noted for its rich cultural and ritualistic life. The construction of large megalithic monuments, such as dolmens and menhirs, points to sophisticated ceremonial practices possibly related to ancestor worship or cosmology. Art from this period includes pottery with intricate designs and engravings on rock surfaces, hinting at symbolic or narrative expressions.

Challenges and Declines

The end of the Chalcolithic period saw several challenges that may have precipitated changes leading into the Bronze Age. Environmental changes, resource depletion, and social tensions within increasingly stratified societies could have led to competition and conflict, catalyzing further societal transformations.

Conclusion

The Iberian Chalcolithic era was a vibrant and dynamic period marked by significant advancements that would critically shape the trajectory of prehistoric Iberia. Its legacy is seen in the technological innovations, social complexity, and artistic expressions that laid a foundation for the subsequent cultural blossoming in the Bronze Age. This period remains a fascinating field of study for archaeologists and historians, offering insights into early European societies' adaptability and ingenuity.

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I2368 Late Chalcolithic Baden Culture, Hungary 3300 BCE Budakalász-Luppa csárda, Hungary View
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I2378 Middle Neolithic Alföld Linear Pottery Culture Tiszadob Group, Hungary 5300 BCE Hejőkürt-Lidl logisztikai központ, Hungary View
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I2022 Early Neolithic Linear Pottery Culture, Germany 5500 BCE Halberstadt-Sonntagsfeld, Germany View
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I2030 Early Neolithic Linear Pottery Culture, Germany 5500 BCE Halberstadt-Sonntagsfeld, Germany View
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I2036 Early Neolithic Linear Pottery Culture, Germany 5500 BCE Halberstadt-Sonntagsfeld, Germany View
I0057 Early Neolithic Linear Pottery Culture, Germany 5215 BCE Halberstadt-Sonntagsfeld, Germany View
I2038 Early Neolithic Linear Pottery Culture, Germany 5350 BCE Halberstadt-Sonntagsfeld, Germany View
I2032 Early Neolithic Linear Pottery Culture, Germany 5500 BCE Halberstadt-Sonntagsfeld, Germany View
I2029 Early Neolithic Linear Pottery Culture, Germany 5299 BCE Halberstadt-Sonntagsfeld, Germany View
I2037 Early Neolithic Linear Pottery Culture, Germany 5210 BCE Halberstadt-Sonntagsfeld, Germany View
I2026 Early Neolithic Linear Pottery Culture, Germany 5500 BCE Halberstadt-Sonntagsfeld, Germany View
I2008 Early Neolithic Linear Pottery Culture, Germany 5211 BCE Halberstadt-Sonntagsfeld, Germany View
I2016 Early Neolithic Linear Pottery Culture, Germany 5500 BCE Halberstadt-Sonntagsfeld, Germany View
I2017 Early Neolithic Linear Pottery Culture, Germany 5500 BCE Halberstadt-Sonntagsfeld, Germany View
I2021 Early Neolithic Linear Pottery Culture, Germany 5500 BCE Halberstadt-Sonntagsfeld, Germany View
I2005 Early Neolithic Linear Pottery Culture, Germany 5299 BCE Halberstadt-Sonntagsfeld, Germany View
I0447 5000 BCE Hódmezővásárhely-Gorzsa. grave 4, Hungary View
I2373 6000 BCE Törökszentmiklos Tiszapüspöki Karanycs haromag 3. lh., Hungary View
I1877 5713 BCE Alsonyek-Bataszek. Mérnöki telep, Hungary View
I3535 5217 BCE Hajdúnánás-Eszlári út, Hungary View
I3536 5300 BCE Enese elkerülő. Kóny. Proletár-dülö. M85. Site 2, Hungary View
I3537 5479 BCE Mezőkövesd-Mocsolyás, Hungary View
I4188 5300 BCE Polgár-Piócás, Hungary View
I0018 5307 BCE Baden-Württemberg. Stuttgart-Mühlhausen. Viesenhaeuser Hof, Germany View
I2739 5307 BCE Alsónyék-Bátaszék. site 11, Hungary View
I2790 3768 BCE Abony. Turjányos-dűlő, Hungary View
I2791 3649 BCE Abony. Turjányos-dűlő, Hungary View
I2793 4442 BCE Törökszentmiklós. road 4. site 3, Hungary View
I1895 5318 BCE Szederkény-Kukorica-dülö, Hungary View
I1902 4900 BCE Felsőörs-Bárókert, Hungary View
I1905 4834 BCE Csabdi-Télizöldes, Hungary View
I1907 4334 BCE Enese elkerülő. Kóny. Proletár-dülö. M85. Site 2, Hungary View
I1878 5830 BCE Alsonyek-Bataszek. Mérnöki telep, Hungary View
I2366 3340 BCE Budakalász-Luppa csárda, Hungary View
I2367 3333 BCE Budakalász-Luppa csárda, Hungary View
I2384 5305 BCE Hajdúnánás-Eszlári út, Hungary View
I0046 5211 BCE Halberstadt-Sonntagsfeld, Germany View
I0659 5210 BCE Halberstadt-Sonntagsfeld, Germany View
I0048 5210 BCE Halberstadt-Sonntagsfeld, Germany View
I1550 5500 BCE Halberstadt-Sonntagsfeld, Germany View
I0821 5205 BCE Halberstadt-Sonntagsfeld, Germany View
I0057 5215 BCE Halberstadt-Sonntagsfeld, Germany View
I2005 5299 BCE Halberstadt-Sonntagsfeld, Germany View
I2008 5211 BCE Halberstadt-Sonntagsfeld, Germany View
I2016 5500 BCE Halberstadt-Sonntagsfeld, Germany View
I2017 5500 BCE Halberstadt-Sonntagsfeld, Germany View
I2021 5500 BCE Halberstadt-Sonntagsfeld, Germany View
I2026 5500 BCE Halberstadt-Sonntagsfeld, Germany View
I2029 5299 BCE Halberstadt-Sonntagsfeld, Germany View
I2030 5500 BCE Halberstadt-Sonntagsfeld, Germany View
I2032 5500 BCE Halberstadt-Sonntagsfeld, Germany View
I2036 5500 BCE Halberstadt-Sonntagsfeld, Germany View
I2037 5210 BCE Halberstadt-Sonntagsfeld, Germany View
I2038 5350 BCE Halberstadt-Sonntagsfeld, Germany View
I3273 3606 BCE Basque Country. Alava. La Chabola de la Hechicera, Spain View
I3276 3096 BCE Basque Country. Alava. Alto de la Huesera, Spain View
I2357 5300 BCE Pusztataskony Ledence 1, Hungary View
I2373 6000 BCE Törökszentmiklos Tiszapüspöki Karanycs haromag 3. lh., Hungary View
Sample ID Culture/Period Date Location Action
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I1302 Chalcolithic Spain 2900 BCE Burgos. Atapuerca. El Mirador Cave, Spain View
I1276 Chalcolithic Spain 2900 BCE Burgos. Atapuerca. El Mirador Cave, Spain View
I1284 Chalcolithic Spain 2900 BCE Burgos. Atapuerca. El Mirador Cave, Spain View
I1274 Chalcolithic Spain 2900 BCE Burgos. Atapuerca. El Mirador Cave, Spain View
I1280 Chalcolithic Spain 2900 BCE Burgos. Atapuerca. El Mirador Cave, Spain View
I1314 Chalcolithic Spain 2900 BCE Burgos. Atapuerca. El Mirador Cave, Spain View
I1277 Chalcolithic Spain 2570 BCE Burgos. Atapuerca. El Mirador Cave, Spain View
I1272 Chalcolithic Spain 2857 BCE Burgos. Atapuerca. El Mirador Cave, Spain View
I1281 Chalcolithic Spain 2867 BCE Burgos. Atapuerca. El Mirador Cave, Spain View
I1300 Chalcolithic Spain 2900 BCE Burgos. Atapuerca. El Mirador Cave, Spain View
I0462 Chalcolithic Spain 2569 BCE Burgos. Arroyal I, Spain View
I0460 Chalcolithic Spain 2460 BCE Burgos. Arroyal I, Spain View
I0459 Chalcolithic Spain 2600 BCE Burgos. Arroyal I, Spain View
I0457 Chalcolithic Spain 3000 BCE Murcia. Caravaca. Camino del Molino, Spain View
I0456 Chalcolithic Spain 3000 BCE Murcia. Caravaca. Camino del Molino, Spain View
I0455 Chalcolithic Spain 2905 BCE Murcia. Caravaca. Camino del Molino, Spain View
I0263 Chalcolithic Spain 2850 BCE Barcelona. Cerdanyola. Paris Street, Spain View
I1553 Chalcolithic Spain 2850 BCE Barcelona. Cerdanyola. Paris Street, Spain View
I0825 Chalcolithic Spain 2471 BCE Barcelona. Cerdanyola. Paris Street, Spain View
I0260 Chalcolithic Spain 2850 BCE Barcelona. Cerdanyola. Paris Street, Spain View
I0261 Chalcolithic Spain 2850 BCE Barcelona. Cerdanyola. Paris Street, Spain View
I0823 Chalcolithic Spain 2850 BCE Barcelona. Cerdanyola. Paris Street, Spain View
I0258 Chalcolithic Spain 2850 BCE Barcelona. Cerdanyola. Paris Street, Spain View
I0262 Chalcolithic Spain 2850 BCE Barcelona. Cerdanyola. Paris Street, Spain View
I0257 Chalcolithic Spain 2573 BCE Barcelona. Cerdanyola. Paris Street, Spain View
I0826 Chalcolithic Spain 2835 BCE Barcelona. Cerdanyola. Paris Street, Spain View
I0458 Chalcolithic Spain 2456 BCE Burgos. Arroyal I, Spain View
I1271 Chalcolithic Spain 2900 BCE Burgos. Atapuerca. El Mirador Cave, Spain View
I1303 Chalcolithic Spain 2900 BCE Burgos. Atapuerca. El Mirador Cave, Spain View
I1981 Chalcolithic Spain 3013 BCE Basque Country. Alava. Alto de la Huesera, Spain View
I0461 Chalcolithic Spain 2453 BCE Burgos. Arroyal I, Spain View
I3270 Chalcolithic Spain 3000 BCE Basque Country. Alava. Las Yurdinas II, Spain View
I3271 Chalcolithic Spain 3000 BCE Basque Country. Alava. Las Yurdinas II, Spain View
I3272 Chalcolithic Spain 3307 BCE Basque Country. Alava. La Chabola de la Hechicera, Spain View
I3269 Chalcolithic Spain 3000 BCE Basque Country. Alava. Las Yurdinas II, Spain View
I4247 Chalcolithic Spain 2462 BCE Madrid. San Fernando de Henares. Camino de las Yeseras, Spain View
I1838 Chalcolithic Spain 3356 BCE Basque Country. Alava. Las Yurdinas II, Spain View
I1843 Chalcolithic Spain 3015 BCE Basque Country. Alava. Alto de la Huesera, Spain View
I1975 Chalcolithic Spain 3092 BCE Basque Country. Alava. La Chabola de la Hechicera, Spain View
I4565 Chalcolithic Spain 3022 BCE Tarragona. Galls Carboners, Spain View
I5838 Chalcolithic Spain 2900 BCE Burgos. Atapuerca. El Mirador Cave, Spain View
I3238 Chalcolithic Spain 2500 BCE Asturias. Soto de las Regueras. Cueva de la Paloma, Spain View
I5665 Chalcolithic Spain 2283 BCE Ciempozuelos. Burgos. Virgazal, Spain View
I6475 Chalcolithic Spain 2500 BCE La Magdalena, Spain View
I6539 Chalcolithic Spain 2500 BCE Madrid. Humanejos, Spain View
I6543 Chalcolithic Spain 2600 BCE Madrid. San Fernando de Henares. Camino de las Yeseras, Spain View
I6584 Chalcolithic Spain 2500 BCE Madrid. Humanejos, Spain View
I6585 Chalcolithic Spain 2397 BCE Madrid. Humanejos, Spain View
I6587 Chalcolithic Spain 2500 BCE Madrid. Humanejos, Spain View
I6588 Chalcolithic Spain 2500 BCE Madrid. Humanejos, Spain View
I6604 Chalcolithic Spain 2130 BCE Madrid. San Fernando de Henares. Camino de las Yeseras, Spain View
I6605 Chalcolithic Spain 2474 BCE Madrid. San Fernando de Henares. Camino de las Yeseras, Spain View
I6610 Chalcolithic Spain 2008 BCE Madrid. San Fernando de Henares. Camino de las Yeseras, Spain View
I6612 Chalcolithic Spain 2600 BCE Madrid. San Fernando de Henares. Camino de las Yeseras, Spain View
I6613 Chalcolithic Spain 2600 BCE Madrid. San Fernando de Henares. Camino de las Yeseras, Spain View
I6617 Chalcolithic Spain 2900 BCE Madrid. Humanejos, Spain View
I6623 Chalcolithic Spain 2600 BCE Madrid. San Fernando de Henares. Camino de las Yeseras, Spain View
I6628 Chalcolithic Spain 2900 BCE Madrid. Humanejos, Spain View
I6629 Chalcolithic Spain 2900 BCE Madrid. Humanejos, Spain View
I6630 Chalcolithic Spain 2900 BCE Madrid. Humanejos, Spain View
I4245 Chalcolithic Spain 2461 BCE Madrid. San Fernando de Henares. Camino de las Yeseras, Spain View
I6472 Chalcolithic Spain 2500 BCE La Magdalena, Spain View
I6609 Chalcolithic Spain 2600 BCE Madrid. San Fernando de Henares. Camino de las Yeseras, Spain View
I3243 Chalcolithic Spain 2500 BCE Asturias. Soto de las Regueras. Cueva de la Paloma, Spain View
I1842 Chalcolithic Spain 3000 BCE Basque Country. Araba/Álava. Peñacerrada-Urizaharra. Las Yurdinas II, Spain View
I1845 Chalcolithic Spain 3100 BCE Basque Country. Araba/Álava. Laguardia. Alto de la Huesera, Spain View
I1846 Chalcolithic Spain 3100 BCE Basque Country. Araba/Álava. Laguardia. Alto de la Huesera, Spain View
I1978 Chalcolithic Spain 3100 BCE Basque Country. Araba/Álava. Laguardia. Alto de la Huesera, Spain View
I7588 Chalcolithic Spain 2900 BCE Andalusia. Córdoba. Lucena. Sima del Ángel, Spain View
I7587 Chalcolithic Spain 2900 BCE Andalusia. Córdoba. Lucena. Sima del Ángel, Spain View
I8048 Chalcolithic Spain 2200 BCE Granada. Arenas del Rey, Spain View
I8132 Chalcolithic Spain 2600 BCE Valencian Community. València/Valencia. La Vital. Gandia, Spain View
I8141 Chalcolithic Spain 2200 BCE Granada. Arenas del Rey, Spain View
I8148 Chalcolithic Spain 2900 BCE Andalusia. Córdoba. Lucena. Sima del Ángel, Spain View
I8149 Chalcolithic Spain 2900 BCE Andalusia. Córdoba. Lucena. Sima del Ángel, Spain View
I8150 Chalcolithic Spain 2900 BCE Andalusia. Córdoba. Lucena. Sima del Ángel, Spain View
I8153 Chalcolithic Spain 2900 BCE Andalusia. Córdoba. Lucena. Sima del Ángel, Spain View
I8154 Chalcolithic Spain 2900 BCE Andalusia. Córdoba. Lucena. Sima del Ángel, Spain View
I8156 Chalcolithic Spain 2900 BCE Andalusia. Córdoba. Lucena. Sima del Ángel, Spain View
I8158 Chalcolithic Spain 2900 BCE Andalusia. Córdoba. Lucena. Sima del Ángel, Spain View
I8197 Chalcolithic Spain 2900 BCE Andalusia. Córdoba. Lucena. Sima del Ángel, Spain View
I8198 Chalcolithic Spain 2900 BCE Andalusia. Córdoba. Lucena. Sima del Ángel, Spain View
I8199 Chalcolithic Spain 2900 BCE Andalusia. Córdoba. Lucena. Sima del Ángel, Spain View
I8364 Chalcolithic Spain 2706 BCE Andalusia. Córdoba. Lucena. Sima del Ángel, Spain View
I8365 Chalcolithic Spain 2706 BCE Andalusia. Córdoba. Lucena. Sima del Ángel, Spain View
I8566 Chalcolithic Spain 3800 BCE Valencian Community. València/Valencia. La Font de la Figuera. Cova de Sant Gomengo, Spain View
I8569 Chalcolithic Spain 2872 BCE Valencian Community. Castelló/Castellón. Alcalá de Xivert. Cova dels Diablets, Spain View
I1976 Chalcolithic Spain 2571 BCE Basque Country. Araba/Álava. Laguardia. El Sotillo, Spain View
I2473 Chalcolithic Spain 2916 BCE Basque Country. Araba/Álava. Laguardia. El Sotillo, Spain View
I3239 Chalcolithic Spain 2500 BCE Asturias. Soto de las Regueras. Cueva de la Paloma, Spain View
I3277 Chalcolithic Spain 3150 BCE Basque Country. Alava. Alto de la Huesera, Spain View
I6622 Chalcolithic Spain 2600 BCE Madrid. San Fernando de Henares. Camino de las Yeseras, Spain View
I8131 Chalcolithic Spain 2576 BCE Valencian Community. València/Valencia. La Vital. Gandia, Spain View
EHU001 Chalcolithic Spain 2287 BCE Castilla y León. Burgos. Monasterio de Rodilla. El Hundido, Spain View
EHU002 Chalcolithic Spain 2564 BCE Castilla y León. Burgos. Monasterio de Rodilla. El Hundido, Spain View
atp016 Chalcolithic Spain 3265 BCE Atapuerca. El Portalón, Spain View
atp002 Chalcolithic Spain 2900 BCE Atapuerca. El Portalón, Spain View
atp12-1420 Chalcolithic Spain 3011 BCE Atapuerca. El Portalón, Spain View
I3276 Chalcolithic Spain 3096 BCE Basque Country. Alava. Alto de la Huesera, Spain View
por002 Chalcolithic Spain 3336 BCE Atapuerca. El Portalón, Spain View
I6542 Chalcolithic Spain 2600 BCE Madrid. San Fernando de Henares. Camino de las Yeseras, Spain View
I0581 Chalcolithic Spain 2900 BCE Burgos. Atapuerca. El Mirador Cave, Spain View
por004 Chalcolithic Spain 3096 BCE Atapuerca. El Portalón, Spain View
I6608 Chalcolithic Spain 2025 BCE Madrid. San Fernando de Henares. Camino de las Yeseras, Spain View
ATP7 Chalcolithic Spain 3345 BCE El Portalon Cave. Sierra de Atapuerca, Spain View
ATP17 Chalcolithic Spain 3008 BCE El Portalon Cave. Sierra de Atapuerca, Spain View
I6596 Chalcolithic Spain 2900 BCE Madrid. Humanejos, Spain View
ATP3 Chalcolithic Spain 3516 BCE El Portalon Cave. Sierra de Atapuerca, Spain View
I6471 Chalcolithic Spain 2500 BCE La Magdalena, Spain View
I8140 Chalcolithic Spain 2121 BCE Granada. Monachil, Spain View
ATP20 Chalcolithic Spain 2291 BCE El Portalon Cave. Sierra de Atapuerca, Spain View
I8155 Chalcolithic Spain 2900 BCE Andalusia. Córdoba. Lucena. Sima del Ángel, Spain View
I6626 Chalcolithic Spain 2600 BCE Madrid. San Fernando de Henares. Camino de las Yeseras, Spain View
I6589 Chalcolithic Spain 2500 BCE Madrid. Humanejos, Spain View
I8157 Chalcolithic Spain 2900 BCE Andalusia. Córdoba. Lucena. Sima del Ángel, Spain View
Ancient Genetic Admixture

Ancient genetic admixture analysis compares the DNA profile of this individual (I3269) with present-day reference populations. These results show what percentage of the individual's genetic makeup resembles ancient populations from different geographic regions.

Neolithic Farmers 55%
European Hunter-Gatherers 23%
Western Steppe Pastoralists 22%
Modern Genetic Admixture

Modern genetic admixture analysis compares the DNA profile of this individual (I3269) with present-day reference populations. These results show what percentage of the individual's genetic makeup resembles modern populations from different geographic regions.

These results complement the ancient ancestry components shown in the previous section, offering a different perspective on the individual's genetic profile by comparing it with modern reference populations rather than prehistoric ancestral groups.

Europe 100%
Southern European 100%
Iberian 51.4%
Sardinian 30.2%
Italian 18.4%
G25 Coordinates

The G25 coordinates for the sample I3269 are as follows. You can analyze its admixture using G25 Studio.

I3269,0.12053822,0.16010046,0.03354468,-0.02705868,0.05716966,-0.0145359,-0.00180504,0.00629234,0.03648532,0.06185382,-0.00592034,0.0144101,-0.0254263,-0.01233356,-0.0034863,0.00356872,0.00307906,-0.00086424,-0.0010269,-0.00314984,0.002313,0.00543502,-0.0120393,-0.0175554,0.00415665
Analyze it in G25 Studio
Scientific Papers References
Parallel palaeogenomic transects reveal complex genetic history of early European farmers
Authors:
Lipson M, Szécsényi-Nagy A, Mallick S
Abstract:

Ancient DNA studies have established that Neolithic European populations were descended from Anatolian migrants who received a limited amount of admixture from resident hunter-gatherers. Many open questions remain, however, about the spatial and temporal dynamics of population interactions and admixture during the Neolithic period. Here we investigate the population dynamics of Neolithization across Europe using a high-resolution genome-wide ancient DNA dataset with a total of 180 samples, of which 130 are newly reported here, from the Neolithic and Chalcolithic periods of Hungary (6000-2900 bc, n = 100), Germany (5500-3000 bc, n = 42) and Spain (5500-2200 bc, n = 38). We find that genetic diversity was shaped predominantly by local processes, with varied sources and proportions of hunter-gatherer ancestry among the three regions and through time. Admixture between groups with different ancestry profiles was pervasive and resulted in observable population transformation across almost all cultural transitions. Our results shed new light on the ways in which gene flow reshaped European populations throughout the Neolithic period and demonstrate the potential of time-series-based sampling and modelling approaches to elucidate multiple dimensions of historical population interactions.

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