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

The Middle to Upper Palaeolithic transition in Europe is associated with the regional disappearance of Neanderthals and the spread of Homo sapiens. Late Neanderthals persisted in western Europe several millennia after the occurrence of H. sapiens in eastern Europe1. Local hybridization between the two groups occurred2, but not on all occasions3. Archaeological evidence also indicates the presence of several technocomplexes during this transition, complicating our understanding and the association of behavioural adaptations with specific hominin groups4. One such technocomplex for which the makers are unknown is the Lincombian-Ranisian-Jerzmanowician (LRJ), which has been described in northwestern and central Europe5-8. Here we present the morphological and proteomic taxonomic identification, mitochondrial DNA analysis and direct radiocarbon dating of human remains directly associated with an LRJ assemblage at the site Ilsenhöhle in Ranis (Germany). These human remains are among the earliest directly dated Upper Palaeolithic H. sapiens remains in Eurasia. We show that early H. sapiens associated with the LRJ were present in central and northwestern Europe long before the extinction of late Neanderthals in southwestern Europe. Our results strengthen the notion of a patchwork of distinct human populations and technocomplexes present in Europe during this transitional period.

Parallel palaeogenomic transects reveal complex genetic history of early European farmers.

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.

The ecology, subsistence and diet of ~45,000-year-old Homo sapiens at Ilsenhöhle in Ranis, Germany.

Recent excavations at Ranis (Germany) identified an early dispersal of Homo sapiens into the higher latitudes of Europe by 45,000 years ago. Here we integrate results from zooarchaeology, palaeoproteomics, sediment DNA and stable isotopes to characterize the ecology, subsistence and diet of these early H. sapiens. We assessed all bone remains (n = 1,754) from the 2016-2022 excavations through morphology (n = 1,218) or palaeoproteomics (zooarchaeology by mass spectrometry (n = 536) and species by proteome investigation (n = 212)). Dominant taxa include reindeer, cave bear, woolly rhinoceros and horse, indicating cold climatic conditions. Numerous carnivore modifications, alongside sparse cut-marked and burnt bones, illustrate a predominant use of the site by hibernating cave bears and denning hyaenas, coupled with a fluctuating human presence. Faunal diversity and high carnivore input were further supported by ancient mammalian DNA recovered from 26 sediment samples. Bulk collagen carbon and nitrogen stable isotope data from 52 animal and 10 human remains confirm a cold steppe/tundra setting and indicate a homogenous human diet based on large terrestrial mammals. This lower-density archaeological signature matches other Lincombian-Ranisian-Jerzmanowician sites and is best explained by expedient visits of short duration by small, mobile groups of pioneer H. sapiens.

Stable isotopes show Homo sapiens dispersed into cold steppes ~45,000 years ago at Ilsenhöhle in Ranis, Germany.

The spread of Homo sapiens into new habitats across Eurasia ~45,000 years ago and the concurrent disappearance of Neanderthals represents a critical evolutionary turnover in our species' history. 'Transitional' technocomplexes, such as the Lincombian-Ranisian-Jerzmanowician (LRJ), characterize the European record during this period but their makers and evolutionary significance have long remained unclear. New evidence from Ilsenhöhle in Ranis, Germany, now provides a secure connection of the LRJ to H. sapiens remains dated to ~45,000 years ago, making it one of the earliest forays of our species to central Europe. Using many stable isotope records of climate produced from 16 serially sampled equid teeth spanning ~12,500 years of LRJ and Upper Palaeolithic human occupation at Ranis, we review the ability of early humans to adapt to different climate and habitat conditions. Results show that cold climates prevailed across LRJ occupations, with a temperature decrease culminating in a pronounced cold excursion at ~45,000-43,000 cal BP. Directly dated H. sapiens remains confirm that humans used the site even during this very cold phase. Together with recent evidence from the Initial Upper Palaeolithic, this demonstrates that humans operated in severe cold conditions during many distinct early dispersals into Europe and suggests pronounced adaptability.

Western visitors at the Blätterhöhle (city of Hagen, southern Westphalia) during the Younger Dryas? A new final palaeolithic assemblage type in western Germany.

Until now, it was considered certain that the last reindeer hunters of the Ahrensburgian (tanged point groups) existed exclusively in northwestern Central Europe during the Younger Dryas Cold Period (~ Greenland Stadial 1). The excavations carried out since 2006 on the forecourt (Vorplatz) of the small Blätterhöhle in Hagen on the northern edge of the Sauerland uplands of southern Westphalia (North Rhine-Westphalia, western Germany) have now changed this view. Beneath a surprisingly extensive sequence of Mesolithic find horizons, Pleistocene sediments could be reached whose excavations yielded a Final Palaeolithic lithic ensemble of the Younger Dryas, unusual for the region and beyond. It is characterised by numerous backed lithic projectile points of high variability. Comparisons suggest a typological-technological connection with the Western European Laborian / Late Laborian. Neither in the nearer nor in the wider surroundings has a comparable lithic find ensemble been found so far. In addition, there is a lack of clear evidence for the reindeer in the fauna. Surprisingly, the vast majority of radiocarbon dates of bones and charcoals from the investigated archaeological horizon of the Final Pleistocene proved to be significantly older than expected from their stratigraphic position. This phenomenon has not yet been clarified.

Low Prevalence of Lactase Persistence in Bronze Age Europe Indicates Ongoing Strong Selection over the Last 3,000 Years.

Lactase persistence (LP), the continued expression of lactase into adulthood, is the most strongly selected single gene trait over the last 10,000 years in multiple human populations. It has been posited that the primary allele causing LP among Eurasians, rs4988235-A [1], only rose to appreciable frequencies during the Bronze and Iron Ages [2, 3], long after humans started consuming milk from domesticated animals. This rapid rise has been attributed to an influx of people from the Pontic-Caspian steppe that began around 5,000 years ago [4, 5]. We investigate the spatiotemporal spread of LP through an analysis of 14 warriors from the Tollense Bronze Age battlefield in northern Germany (∼3,200 before present, BP), the oldest large-scale conflict site north of the Alps. Genetic data indicate that these individuals represent a single unstructured Central/Northern European population. We complemented these data with genotypes of 18 individuals from the Bronze Age site Mokrin in Serbia (∼4,100 to ∼3,700 BP) and 37 individuals from Eastern Europe and the Pontic-Caspian Steppe region, predating both Bronze Age sites (∼5,980 to ∼3,980 BP). We infer low LP in all three regions, i.e., in northern Germany and South-eastern and Eastern Europe, suggesting that the surge of rs4988235 in Central and Northern Europe was unlikely caused by Steppe expansions. We estimate a selection coefficient of 0.06 and conclude that the selection was ongoing in various parts of Europe over the last 3,000 years.

The Late Upper Palaeolithic and earliest Mesolithic evidence of burials in Europe.

Burials of the Late Palaeolithic (14 000-11 600 cal years before present, henceforth BP) are a rare phenomenon in Europe. Several sites possess burials of single and double individuals. As with the preceding Magdalenian, the burial of more than two individuals in the same grave cutting seems to be unusual, but does occur occasionally. The deposition of isolated and disarticulated human remains with or without cut marks seems additionally to belong to the Magdalenian context. In the final Palaeolithic phase (13 000-11 600 cal years BP) there is evidence for cemetery-like clusters of burials, which contrast to the Magdalenian evidence, instead showing some similarities with the succeeding Mesolithic. The earliest Mesolithic burials 11 600-10 500 cal BP) are a very rare phenomenon, covering a short time span between the beginning of the Preboreal and the beginning of the Boreal phase of the early Holocene. Here the evidence includes single inhumations, cemetery-like structures and a number of isolated human remains. Caves and rock shelters were the most common places for inhumations in both the final Palaeolithic and the early Mesolithic. Although the number of sites with a chronological continuity from the LUP to the Early Mesolithic burial is low, several aspects indicate a general continuity in burial patterns over this period. Apart from this continuity, the Mesolithic burials in general seem to represent a new level of diversity in burial practices.This article is part of the theme issue 'Evolutionary thanatology: impacts of the dead on the living in humans and other animals'.

Flint arrowhead embedded in a human humerus from the Bronze Age site in the Tollense valley, Germany - A high-resolution micro-CT study to distinguish antemortem from perimortem projectile trauma to bone.

The Bronze Age site in the Tollense valley, Germany, has yielded thousands of human and animal bones and a number of archaeological artifacts. Several of the human bones exhibit blunt and sharp force lesions, and the assemblage has been interpreted as representing victims of a large scale conflict. One of the earliest finds is a human humerus with an embedded flint arrowhead. Alleged signs of healing initially reported for this humerus based on clinical CT imaging were interpreted as evidence of an antemortem lesion. The present study, using micro-CT imaging, revealed that the arrowhead lesion in the humerus, contrary to the previous interpretation, shows no signs of healing. The structure previously assumed to represent a sclerotic margin around the wound canal was shown to actually represent compacted trabecular debris. Thus, our re-analysis of the specimen led to a re-classification of the arrow wound as a perimortem lesion. The findings of the present study demonstrate the value of micro-CT imaging as a non-destructive method for obtaining information on the nature of bone lesions and healing reactions critical for the reconstruction of interpersonal conflict scenarios in the past.

2000 years of parallel societies in Stone Age Central Europe.

Debate on the ancestry of Europeans centers on the interplay between Mesolithic foragers and Neolithic farmers. Foragers are generally believed to have disappeared shortly after the arrival of agriculture. To investigate the relation between foragers and farmers, we examined Mesolithic and Neolithic samples from the Blätterhöhle site. Mesolithic mitochondrial DNA sequences were typical of European foragers, whereas the Neolithic sample included additional lineages that are associated with early farmers. However, isotope analyses separate the Neolithic sample into two groups: one with an agriculturalist diet and one with a forager and freshwater fish diet, the latter carrying mitochondrial DNA sequences typical of Mesolithic hunter-gatherers. This indicates that the descendants of Mesolithic people maintained a foraging lifestyle in Central Europe for more than 2000 years after the arrival of farming societies.

A critical review of the German Paleolithic hominin record.

We review the hominin fossil record from western Central Europe in light of the recent major revisions of the geochronological context. The mandible from Mauer (Homo heidelbergensis), dated to circa 500,000 years ago, continues to represent the earliest German hominin and may coincide with the occupation of Europe north of the high alpine mountain chains. Only limited new evidence is available for the Middle Pleistocene, mostly in the form of skull fragments, a pattern that may relate to taphonomic processes. These finds and their ages suggest the gradual evolution of a suite of Neandertal features during this period. Despite new finds of classic Neandertals, there is no clear proof for Neandertal burial from Germany. Alternatively, cut marks on a skull fragment from the Neandertal type site suggest special treatment of that individual. New Accelerator Mass Spectrometry (AMS) radiocarbon dates of previous finds leave little reliably dated evidence for anatomically modern humans (AMH) in Europe before 30,000 BP; the remains from Hahnöfersand, Binshof-Speyer, Paderborn-Sande, and Vogelherd are now of Holocene age. Thus, a correlation of AMH with the Aurignacian remains to be proven, and the general idea of a long coexistence of Neandertals and AMH in Europe may be questioned. In western Central Europe, evidence of Gravettian human fossils is also very limited, although a new double grave from lower Austria may be relevant. The only dated burial from the German Upper Paleolithic (from Mittlere Klause) falls into a time period (circa 18,600 BP) represented by only a few occupation sites in western Central Europe. A number of human remains at Magdalenian sites appear to result from variable (secondary) burial practices. In contrast, the Final Paleolithic (circa 12,000-9600 cal. BC) yields an increase of hominin finds, including multiple burials (Bonn-Oberkassel, Neuwied-Irlich), similar to the situation in western and southern Europe.