Homo sapiens reached the higher latitudes of Europe by 45,000 years ago

The study investigates who produced the Lincombian-Ranisian-Jerzmanowician (LRJ) technocomplex during the Middle to Upper Palaeolithic transition and clarifies the timing and geographic extent of early Homo sapiens in Europe. The LRJ, distributed across northwestern and central Europe, has been variously attributed to Neanderthals or H. sapiens due to its mix of laminar blade production, partial-bifacial blade points (Jerzmanowice points), and occasional bifacial leaf points. Chronologically, LRJ assemblages (~44,000–41,000 cal BP) overlap with the presence of both Neanderthals and H. sapiens in Europe, making attribution uncertain. The site Ilsenhöhle in Ranis, Germany—an eponymous LRJ locality—provides stratified contexts and materials to test whether LRJ assemblages were made by H. sapiens, and to place this in the broader framework of the Neanderthal–modern human transition, population interactions, and behavioral adaptations under variable MIS 3 climatic conditions.

Prior research documents that late Neanderthals persisted in parts of western Europe after H. sapiens appeared in the east, with evidence for local hybridization but not universally. The LRJ has been described across the British Isles, northwestern, and central Europe and debated as either Early Upper Palaeolithic linked to H. sapiens or as a local Neanderthal development due to its bidirectional blade production and presence of bifacial elements suggesting Middle Palaeolithic roots. LRJ assemblages generally date to ~44,000–41,000 cal BP, overlapping with both populations. Comparative contexts include Initial Upper Palaeolithic sequences like Bacho Kiro Cave (Bulgaria) and Bohunician (Moravia), as well as contemporaneous bifacial industries (Szeletian, Altmühlian). The disputed early H. sapiens evidence at Kent’s Cavern (UK) and Grotte Mandrin (France) further points to a complex mosaic of occupations during 55,000–45,000 cal BP.

- Fieldwork and stratigraphy: Reopened and extended the 1930s excavations at Ilsenhöhle in Ranis in 2016–2022 to bedrock, documenting layers 11–6 with correlations to the historical stratigraphy (layer X ≈ layers 9–8). Identified lithic concentrations, raw material sourcing (Baltic flint; some quartzite), and roof collapse sealing events. - Radiocarbon dating and modeling: Constructed a Bayesian chronological model from 28 radiocarbon dates on material from layers 11–7, including directly dated human remains, modified bones, and charcoal. Collagen yields averaged 11.8% (range 5.3–16.3%, n=33); only one outlier identified, supporting stratigraphic integrity. Directly dated six human bones from the 1930s LRJ layer X collection. Calibrated ranges reported at 95.4% probability. - Proteomic screening and identification: Applied MALDI-TOF MS and LC–MS/MS proteomics, complemented by morphological assessment, to screen bone assemblages from 2016–2022 and 1932–1938 excavations. Identified 13 hominin bone specimens in total (4 from new excavations; 9 from the historical collection). Assessed diagenesis via collagen deamidation and calculated proteomic coverage per amino acid against Homininae references. Proteomics confirmed Homininae but could not resolve finer taxonomic distinctions among hominin populations due to database limitations. - Ancient DNA: Screened 11 hominin remains for mtDNA; recovered 4,413–175,688 unique reads per specimen with characteristic C-to-T damage patterns. Reconstructed near-complete mtDNA genomes for 10 fragments; assigned to H. sapiens using diagnostic positions distinguishing from Neanderthal and Denisovan. Performed phylogenetic analyses using BEAST2 to place mtDNA haplotypes among ancient and modern human references. - Zooarchaeology and paleoenvironment: Identified fauna via zooarchaeology and proteomics (17 taxa) with predominance of reindeer; evidence indicates primary accumulation by large carnivores with sporadic human use. Conducted sedimentology, micromorphology (including charred plant inputs), and stable isotope analyses (equid teeth) to reconstruct climate (temperature declines, open steppe, highly seasonal subarctic conditions). Linked occupations to Greenland stadial/interstadial phases (GS13, GI12, GS12). - Lithic analysis: Documented LRJ blade production (fragmented blades, Jerzmanowice points, bifacial leaf points in the historical collection). Raw material analysis showed links to northern lowlands (Baltic flint).

- Direct association of H. sapiens with LRJ at Ranis: 11 tested skeletal fragments yielded H. sapiens mtDNA; 10 near-complete mtDNA genomes reconstructed. Five genomes showed no pairwise differences (same or maternally related individuals). - Chronology: Layer 11 dates to 55,860–48,710 cal BP. LRJ layers 9 and 8 date to 47,500–45,770 cal BP and 46,820–43,260 cal BP respectively (95.4% probability). Directly dated LRJ-associated human bones from the 1930s collection fall within 46,950–42,200 cal BP (95.4% probability), consistent with layers 9–8. - Haplogroups and phylogeny: 9/10 Ranis mtDNA genomes are haplogroup N; 1/10 is haplogroup R. The N haplotypes cluster with the Zlatý kůň individual; genetic date estimates for Ranis mtDNA range 49,105–40,918 years BP, consistent with radiocarbon results. - Occupation and environment: Lithic densities in LRJ layers are modest; Baltic flint predominates with some quartzite. Zooarchaeology and proteomics identified 17 taxa (dominant reindeer; also aurochs/bison, red deer, horse, woolly rhinoceros, mammoth; carnivores dominated by cave bear). Evidence indicates sporadic, short-term human use amid carnivore accumulation. Stable isotopes indicate a cold open steppe with temperatures ~7–15 °C below modern during ~45,000–43,000 cal BP (overlapping layers 8 and 7), consistent with highly seasonal subarctic climate. - Regional significance: Demonstrates H. sapiens were present in central and northwestern Europe by at least ~45,000 cal BP, preceding the final disappearance of Neanderthals in southwestern Europe (~42,000 cal BP).

The findings resolve a long-standing debate by demonstrating that the LRJ technocomplex at Ranis was produced by Homo sapiens. This establishes that pioneer H. sapiens reached higher mid-latitudes of Europe earlier than previously documented for the Aurignacian, during a time of markedly cold, open steppe environments. The overlap in timing with late Neanderthals in other regions argues for a patchwork of contemporaneous populations and technocomplexes across Europe rather than a simple replacement sequence. MtDNA affinities linking Ranis individuals to Zlatý kůň and, in one case, to Fumane 2, connect LRJ-associated H. sapiens to a broader network of early incursions in central and eastern Europe, potentially relating LRJ to IUP phenomena such as the Bohunician. The evidence challenges the hypothesis of a long hiatus between Neanderthal disappearance and H. sapiens arrival in northwestern Europe and suggests more complex, regionally variable dynamics of dispersal, interaction, and cultural expression during the transition.

This study provides direct evidence that H. sapiens produced the LRJ at Ilsenhöhle in Ranis and occupied higher-latitude regions of Europe by ~45,000 cal BP in cold, highly seasonal conditions. By combining stratified excavation, radiocarbon modeling, proteomics, and ancient mtDNA, the work clarifies the makers of a key transitional technocomplex and situates them within a broader early H. sapiens network in Europe. The results reject a simple hiatus model for northwestern Europe and instead support a mosaic of populations and industries. Future research should: (1) investigate genomic (nuclear DNA) relationships of LRJ-associated individuals to other early European populations; (2) clarify technological and cultural links among LRJ, Bohunician, Szeletian, and Altmühlian industries; (3) refine regional chronologies and site-use patterns; and (4) expand paleoecological reconstructions to understand how climate shaped early H. sapiens dispersal and behavior.

- Proteomic taxonomy is limited by current reference databases and cannot resolve distinctions within Homininae beyond genus-level confirmation. - Ancient DNA results are based on mtDNA; nuclear genome data were not reported, limiting resolution of population structure, admixture, and individual counts. - Some historical collections had mixed layer labels due to 1930s excavation methods, introducing provenience uncertainty (mitigated by new stratigraphic work and direct dating). - Diagnostic LRJ points were not recovered in the new excavations, relying partly on the 1930s assemblage for typological attribution. - Evidence suggests sporadic, short-term human occupations with low artefact densities, which can bias behavioral reconstructions. - Broader regional attributions (e.g., relationship to Bohunician, Szeletian, Altmühlian) remain inferential and require further comparative work.