This is a comprehensive update to the previously proposed extinct maternal haplogroup U10. This lineage is now being officially renamed U11 to avoid confusion with the new, living U10 haplogroup (defined by FTDNA) recently assigned to a minor branch from the South Caucasus. https://discover.familytreedna.com/mtdna/U10/scientific?section=variants
1. Phylogenetic Position and Defining MarkersThe U11 haplogroup represents a distinct, extinct maternal lineage found across Paleolithic and Mesolithic Western Eurasia.
Parent Clade: U2'3'4'7'8'9a
Derivation: U11 is derived from the lineage that shares the defining marker T5999C with the U4'9'10 branches. The macro-clade uniting these is now referred to as U4'9'10'11.
Distinction from FTDNA's U10: Crucially, U11 does not share the G499A marker that defines the U4'9'10 branch leading to the FTDNA-classified U10 (the minor, living branch).
Core Defining Markers of U11: The proposed U11 clade is defined by the unique presence of these four markers: 10020C, 15466A, 6152C, 16297C.
2. Evidence from Ancient Genomes: More than 10 fossils with Western Paleo-Eurasian traits have been found that belong to this maternal haplogroup, which is derived from U2'3'4'7'8'9 and runs parallel to U4'9, sharing the 5999C marker.The analysis of suspicious ancient mitochondrial sequences suggests that 15466A and 6152C are the common ancestral features to all samples of U11.The oldest known sample of this maternal lineage was found in the Malalmuerzo Cave (Granada, Andalusia, Spain, ca. 23,000 BP) and was published in 2023 as two samples from the same individual (MLZ003 and MLZ005). Recently, the complete genome of the child from Grotta delle Mura (Apulia, Italy, ca. 17,000 BP) was published in 2024, confirming the core markers of U11. The markers shared by Malalmuerzo and Grotta delle Mura that define the U11 haplogroup are the four core markers listed above. These four variants are also shared with the nearly contemporaneous sample LMA001 from La Marche (France, 16,200 BP).The systematic renaming to U11 clarifies the distinction between this extinct Western Eurasian Paleolithic/Mesolithic lineage and the recently defined living U10 branch.
3. Key Ancient Samples Confirmed as U11: The following table summarizes confirmed ancient samples belonging to the proposed U11 haplogroup lineage, spanning from the Late Paleolithic through the Mesolithic
Sample ID,Site Location,Approximate Age (cal BP),Key Context & Defining Markers,Parent Clade
MLZ003/005,"Malalmuerzo Cave, Spain","ca. 23,000 BP (Gravettian)",Oldest known sample of the clade. Core U11 markers confirmed.,U4'9'10'11 (5999C)
Mura1,"Grotta delle Mura, Italy","ca. 17,000 BP","Complete genome, confirming core U11 markers (10020C, 15466A, 6152C, 16297C).",U4'9'10'11 (5999C)
LMA001,"La Marche, France","16,273–15,958 BP (Magdalenian)","Core U11 markers + Extras (150T, 152C, 310C, 6498A, 14152G, 16274A, 16519C).",U4'9'10'11 (5999C)
RIP001,"Riparo Tagliente, Italy","15,026–14,560 BP",Core U11 markers + Extras (13183G). Includes the 14152G extra marker reported in other samples.,U4'9'10'11 (5999C)
MAZ003,"Maszycka, Poland","13,804–13,407 BP","Core U11 markers + Extras (14152G, 13183G).",U4'9'10'11 (5999C)
STO001,"San Teodoro, Italy (Sicily)","11,627–11,397 BP",Core U11 markers + 16297C marker is missing.,U4'9'10'11 (5999C)
BAL003,"Balma Guilanyà, Spain","10,727–9,272 BP (Mesolithic)","Shows long-term Iberian presence. Core U11 markers, but 6152C and 16297C are missing.",U4'9'10'11 (5999C)
UZZ096,"Grotta dell'Uzzo, Italy (Sicily)","8,800–7,800 BCE (Mesolithic)","Core U11 markers, but 16297C is missing.",U4'9'10'11 (5999C)
NEO694,"Santa Maira, Spain","7,648–7,496 BP",Core U11 markers + Extras (12245C).,U4'9'10'11 (5999C)
I2158,"Grotta d'Oriente, Italy (Sicily)","ca. 12,350–7,750 BCE",Core U11 markers confirmed.,U4'9'10'11 (5999C)
Recently, more ancient samples of this extinct haplogroup have been found in El Mirón Cave (Asturias, Spain) and Romito (Italy), making it the second most common haplogroup in the Western Mediterranean Arc during the Magdalenian period, second only to U5b.
[UPDATE — March 2026] New Gravettian genomic data from the Franco-Cantabrian region: implications for the phylogenetic continuity debate and the extinct haplogroup U11
Two recent studies by Pere Gelabert and collaborators (University of Vienna / Harvard) substantially advance our understanding of Upper Palaeolithic maternal and nuclear genetic diversity in southwestern Europe, with direct implications for the arguments presented in this blog.
1. Sedimentary ancient DNA at El Mirón: the Solutrean U2'3'4'7'8'9 signal
The published study (Gelabert et al., 2025, Nature Communications, DOI: 10.1038/s41467-024-55740-7) recovers human mitochondrial DNA from Solutrean sediment layers at El Mirón Cave (Cantabria, ~22,000 cal BP). The human consensus sequences from the best-preserved sample (ElMirón14, Solutrean Layer 122) are assigned to haplogroup U2'3'4'7'8'9, confirming and extending the classification previously reported for this site. This basal notation represents an unresolved node at the root of the U2 through U11 subtree — a paraclade that encompasses the extinct haplogroup U11 discussed throughout this blog.
As the authors note, haplogroup U2'3'4'7'8'9 has been identified in only approximately thirteen individuals worldwide to date, all associated with Gravettian, Solutrean, or early Magdalenian contexts, distributed from Poland to the Iberian Peninsula, and restricted to a time window of roughly 27,000 to 13,000 years ago. This temporal and geographic restriction is consistent with U11 — and its sister lineages U9 and U10 — representing derived branches of precisely this Upper Palaeolithic western European clade that failed to survive into the post-LGM world. The El Mirón sedaDNA data thus provide new stratigraphic confirmation of the persistence of this ancestral U-paraclade node within the Franco-Cantabrian refugium during the Last Glacial Maximum itself.
2. Gravettian genomes from Chufín, Isturitz, and Ostuni: new mtDNA haplogroups
The preprint (Gelabert et al., preprint 2025, bioRxiv, DOI: 10.1101/2025.09.26.678744) presents genome-wide data from three new Gravettian individuals:
Chufín Cave (Cantabria, ~22,793 cal BP, female): mtDNA haplogroup U8. This same haplogroup is shared with Dolní Věstonice 13 (Gravettian, Czechia), Goyet Q116-15 (Aurignacian, Belgium), and BK-1653 from Bacho Kiro Cave (Initial Upper Palaeolithic, Bulgaria). Notably, the U8a subclade — a derived branch of U8 — survives to the present day at unusually high frequencies among modern Basques, suggesting a deep regional connection. Chufín itself is geographically adjacent to the Basque Country.
Isturitz Cave (French Basque Country, ~30,850–29,970 cal BP, female): mtDNA haplogroup M. This is a deep basal lineage within macro-haplogroup M, and the same haplogroup carried by both Ostuni1 and her son Ostuni1b from southern Italy. As the authors explicitly state: "Haplogroup M is a common haplogroup in Europe before the LGM, but very rare among post-LGM individuals."
Ostuni1b (southern Italy, ~27,810–27,430 cal BP, male): mtDNA haplogroup M, genetically consistent with being the son of the previously published Ostuni1 individual.
3. Haplogroup M at Isturitz and the Franco-Cantabrian refugium: a case for maternal discontinuity
The haplogroup M assignment at Isturitz is, arguably, the most consequential mitochondrial finding in this preprint for the Franco-Cantabrian refugium debate. The site of Isturitz has long been recognised as a major aggregation hub in the Upper Palaeolithic, located at the critical passage between continental Europe and the Iberian Peninsula at the western end of the Pyrenees, and characterised by an exceptionally rich portable art assemblage with close stylistic parallels across the Pyrenean and Cantabrian regions. Its genomic record now shows that the female buried there ~30,000 years ago carried haplogroup M — a lineage that, within the Iberian Peninsula, has previously been documented only at Catalan Gravettian sites (including Mollet III and Reclau Viver, both in Girona province), based on the dataset of Fu et al. (2016) and Posth et al. (2023).
The phylogenetic implication is straightforward and directly relevant to the continuity debate. If the Franco-Cantabrian refugium had maintained strict maternal genetic continuity from the Gravettian through the Solutrean and into the Magdalenian, haplogroup M should appear — however infrequently — in post-LGM samples from the same region. It does not. No Magdalenian individual from Iberia or southern France has been assigned haplogroup M to date. This absence, combined with the nuclear-level evidence presented in the same preprint — specifically, the f-statistics demonstrating that none of the Cantabrian Solutrean or Gravettian samples show affinity to the GoyetQ2 Magdalenian cluster, and that Villabruna-related ancestry arrives in northern Iberia only after the LGM — constitutes strong evidence against a model of uninterrupted local population transmission.
The picture that emerges is one of partial replacement: a pre-LGM Franco-Cantabrian population carrying diverse maternal lineages (M at Isturitz, U8 at Chufín, U2'3'4'7'8'9 in the Solutrean sediments of El Mirón) that were largely not transmitted to the post-LGM Magdalenian groups. The extinct haplogroup U11, like haplogroup M and like the unresolved U2'3'4'7'8'9 node itself, represents one of those maternal threads that did not survive the demographic bottleneck associated with the Last Glacial Maximum.
References for this update:
Gelabert, P., Oberreiter, V., Straus, L. G., González Morales, M. R., Sawyer, S., Marín-Arroyo, A. B. et al. (2025). A sedimentary ancient DNA perspective on human and carnivore persistence through the Late Pleistocene in El Mirón Cave, Spain. Nature Communications, 16(1), 107. DOI: 10.1038/s41467-024-55740-7.
Gelabert, P., Sawyer, S., Cheronet, O., Villalba-Mouco, V., Oberreiter, V., González-Morales, M. R., Straus, L. G. et al. (preprint, 2025). Genetic admixture between East and West European Gravettian-associated populations in Western Europe before the Last Glacial Maximum. bioRxiv. DOI: 10.1101/2025.09.26.678744.
Fu, Q. et al. (2016). The genetic history of Ice Age Europe. Nature, 534, 200–205. DOI: 10.1038/nature17993.
Posth, C. et al. (2023). Palaeogenomics of Upper Palaeolithic to Neolithic European hunter-gatherers. Nature, 615, 117–126. DOI: 10.1038/s41586-023-05726-0
Additional reference for the bibliography:
Villalba-Mouco, V. et al. (2023). A 23,000-year-old southern Iberian individual links human groups that lived in Western Europe before and after the Last Glacial Maximum. Nature Ecology & Evolution, 7(4), 597–609. DOI: 10.1038/s41559-023-01987-0.
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