Archaeological, osteological and genetic evidence suggests that Neanderthals lived in small groups1,2; however, less is known about whether these groups were part of isolated communities or belonged to larger, well-connected populations3. The dense concentration of broadly contemporaneous Neanderthal sites in the Meuse Basin, Belgium4, provides a rare opportunity to study regional populations at high resolution. Here we generated genetic data from 27 Neanderthals who lived less than approximately 52,500 years ago from ten archaeological sites in Belgium and France, including a high-coverage genome from a 45,000-year-old individual from Goyet, Belgium. We show that most of these individuals are more closely related to one another than to other contemporaneous late Neanderthals in Europe. Further, some of these individuals carry DNA from a Neanderthal lineage predating the split of late Neanderthals. Although these Neanderthals overlapped temporally with early modern humans in northwestern Europe from around 47,000 years ago, we find no evidence of recent gene flow from modern humans. They also do not show the genetic signatures of mating among close relatives found in Altai Neanderthals, suggesting that they lived in larger or better-connected groups. Moreover, genetic load did not accumulate over time, arguing against progressive genetic deterioration as a driver of Neanderthal extinction.
Why does research on Neanderthals attract public attention far beyond its immediate scientific relevance? Such fascination may reflect more than intellectual curiosity; it may involve the activation of deep symbolic structures that Carl Jung termed the collective unconscious. Integrating evolutionary biology, genomics, paleoanthropology, and psychology, I address a question Jung did not explicitly pose: when, along the human evolution, did the collective unconscious originate? I argue that this structure emerged gradually rather than suddenly. Homo erectus established a cognitive floor characterized by basic schemas of fear, group cohesion, and hierarchy, without evidence of symbolic elaboration. Homo heidelbergensis, the common ancestor of Neanderthals and Homo sapiens, is proposed as the strongest candidate for the emergence of proto-archetypal cognition, given its enlarged brain, social complexity, and possible forms of mortuary behavior. The symbolic system was operational in Neanderthals and archaic H. sapiens and became fully visible with the symbolic expression of the Upper Paleolithic. I further introduce the concept of the "incorporated other" to explain the distinctive psychological salience of Neanderthals and Denisovans. Unlike other extinct organisms, these archaic humans remain present in contemporary populations through genetic introgression, collapsing boundaries between self and other, past and present. I propose that this condition contributes to public fascination in ways not fully explained by resemblance, category ambiguity, intuitive essentialism, or the uncanny. The "incorporated other" predicts that genetic incorporation itself increases perceived self-relevance and engagement, even when similarity is held constant. Neanderthals are therefore psychologically salient because they are simultaneously extinct and biologically present.
The southern coast of Europe has been at the center of archaeological debates contrasting the social and cognitive capabilities of Neanderthals and modern humans. Early evidence of marine resource exploitation by Neanderthals in this region challenged some views that coastal adaptation was a trait distinctive of modern human cognition and behavior. While it is now evident that Neanderthals exploited marine resources, the nature of their seasonal foraging strategies remains an open question. Here, we analyzed the oxygen isotope composition of mollusk shells collected by Neanderthals at Los Aviones Cave (Iberian Peninsula), demonstrating that they primarily exploited intertidal mollusks during the colder months of the year. This harvesting pattern anticipates the dominant seasonal exploitation strategies observed among later modern human populations in southern Europe, highlighting a degree of behavioral flexibility in Neanderthal use of coastal ecosystems.
The demographic history of Neanderthals is only partially understood. In Europe, some degree of genetic continuity has been shown from 120 thousand years ago (ka) onward despite the occurrence of multiple subsequent diversification events. While it has been proposed that a population turnover preceded the emergence of Late Neanderthals in Europe, the extent, timing, and geographic location of this event are currently unknown. Here, we report ten mitochondrial DNA sequences (mtDNAs) of Neanderthal individuals from six archaeological sites in Belgium, France, Germany and Serbia, and analyze them alongside 49 published mtDNAs. The integration of phylogenetic and molecular dating analyses with an extensive archaeological dataset enabled us to reconstruct temporal and spatial patterns in Neanderthal distribution. Remarkably, nearly all Late Neanderthal individuals across Europe belong to a single mtDNA lineage that diversified recently, confirming a large-scale genetic replacement. Our analyses date this diversification event to approximately 65 ka and suggest that it likely originated from a population refugium in southwestern France from which Neanderthals appear to have undergone a major range dispersal across Europe. In addition, we detect a sharp decline in the Neanderthal mtDNA effective population size beginning ~45 ka and reaching a minimum ~42 ka, shortly before their extinction. This study demonstrates that integrating molecular and archaeological datasets provides a more detailed understanding of the Late Neanderthal population's history, and highlights the critical role of climate-driven refugia and subsequent range expansions in shaping the genetic landscape of Neanderthals through time.
The use of faunal remains as tools by Neanderthals has long intrigued researchers. These remains include mammal teeth, which are particularly durable and resistant. Nevertheless, there is a significant gap in taphonomic analysis of dental remains, and archaeological experiments with tooth tools remain scarce. Recent studies suggest that Paleolithic groups may have used rhinoceros teeth as tools. This work seeks to elucidate this question by applying a multidisciplinary approach, including taphonomic and traceological analyses using various microscopic imaging techniques. We applied this approach to several rhinoceros teeth from key archaeological, paleontological, and contemporary collections. For the first time, we also set up controlled archaeological experiments on rhinoceros teeth. The results shed light on the potential role of rhinoceros teeth as versatile tools during the Middle Paleolithic in Western Europe and contribute to our understanding of Neanderthals' adaptive subsistence strategies and material culture.
Neanderthal medical knowledge has long attracted scholarly interest. Evidence suggests they cared for sick, injured, and elderly group members, with possible use of medicinal plants. However, it remains uncertain whether such practices reflect deliberate medical strategies or instinctive self-medication akin to that observed in non-human primates. Here, we analyze and interpret traces of deliberate artificial manipulation of Chagyrskaya 64, a Neanderthal lower left second molar found in Chagyrskaya Cave (Altai Krai, Russia). The tooth exhibits a large human-generated concavity on the occlusal surface, created during the lifetime of the individual. Traceological and microtomographic analyses of the observed modifications, combined with experimental verification, reveal that the concavity in Chagyrskaya 64 is indicative of the earliest documented instance of caries treatment involving the drilling/rotating with a lithic perforator, ca. 59 ka. Evidence of two distinct types of manipulations requiring different tools, in addition to the drilling/rotating technique, necessitating complex finger movements, indicates that the Chagyrskaya Cave Neanderthals possessed the cognitive capacity to intuit the source of pain, comprehend the feasibility of its elimination, and deliberately select the most efficacious dental intervention. These patterns bring Neanderthal behavior closer to modern humans and differentiate that behavior from the instinctive actions of other primates.
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[This corrects the article DOI: 10.1371/journal.pone.0347662.].
De Nadale Cave is a Neanderthal site located in the Berici Hills of Northeast Italy that provides important insights into regional Middle Paleolithic occupational dynamics. This paper presents the results of the first combined uranium-series and electron spin resonance (U-S/ESR) dating study of herbivore teeth recovered from the site. Three bovid and cervid teeth yield U-S/ESR ages of 79 ± 9 ka, 72 ± 11 ka, and 55 ± 11 ka, with a calculated weighted mean age of 70.1 ± 5.9 ka. These age estimates align with, and further clarify, earlier biochronological findings at De Nadale Cave, as well as previous analyses of charcoal fragments and U/Th dating undertaken on a large herbivore tooth from the site. The new U-S/ESR ages are statistically indistinguishable from the previously published U/Th minimum age of 70.2 +1.0/-0.9 ka when considering their 2σ uncertainty ranges. The consistency of these results supports the reliability of the U-S/ESR chronological framework at De Nadale and indicates that Neanderthal occupation of the site potentially occurred close to the transition between late Marine Isotope Stage (MIS) 5 and early MIS 4, providing important insights into Neanderthal occupation patterns during the harsh stadial and transitional climate phases of the Late Pleistocene. More broadly, the improved chronological framework for De Nadale indicates that Middle Paleolithic Quina technology is likely associated with this MIS transition, in accordance with emerging chronological findings from similar archaeological sites across Western Europe.
A pair of studies illuminates these humans' long, hardscrabble existence.
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We present evidence from Üçağızlı II Cave in the northern Levant, Türkiye, documenting the sequential occupation by Neanderthals (Homo neanderthalensis; 77-59 ka) and modern humans (Homo sapiens; 59-47 ka). The combined evidence of human fossils, faunal and floral remains, lithics, and manuports demonstrates significant behavioral and technological continuity across this taxonomic sequence, characterized by consistent subsistence strategies and the persistent selection and transport of specific nondietary mollusk shells. The lithic assemblages largely align with late Middle Paleolithic traditions, exhibiting characteristic Mousterian variants. The association of diagnostic human fossils with these archaeological remains demonstrates that the shift from Neanderthals to modern humans occurred within a cultural continuum. Our findings suggest shared behaviors between Neanderthals and modern humans that extended beyond subsistence to include nonutilitarian behaviors within the specific geographic and temporal context studied here.
The site of Lehringen (Germany) has played a pivotal role in the study of the hunting behavior of Neanderthals. The finding of a 2.38 m long wooden thrusting spear was at the time of discovery in 1948 the only complete Palaeolithic hunting tool, dating back to the last interglacial, approximately 125 ka. The interrelation of the thrusting spear with the skeleton of a straight-tusked elephant has both been interpreted as direct evidence of hunting, or as a coincidental association. Here we report on results of the first systematic analysis of the faunal assemblage of the site including the remains of the straight-tusked elephant. Cut marks on several ribs and vertebrae of the elephant show that the animal was defleshed and probably eviscerated in a rather fresh state, indicating early access to the carcass by Neanderthals. Additionally, remains of aurochs, brown bear, and beaver show signs of butchery. It demonstrates that Neanderthals of the last interglacial at the northern limits of their known distribution were exploiting a wide range of animals on different occasions, including the largest prey of that time. In conclusion, Lehringen represents the most convincing Neanderthal site with evidence of a successful elephant hunt with a thrusting spear and demonstrates opportunistic hunting behavior during the Eemian.
Data on palaeolithic subsistence is often obtained through studies of faunal palimpsests, containing remains of animal processing activities accumulated over non-quantifiable amounts of time. Compounding such site-specific data with evidence from other sites distributed over large areas - i.e. integrating data spanning large temporal as well as spatial scales - results in coarse-grained reconstructions of past prey diversity. In contrast, here we present prey diversity data from what is—geologically speaking—a “snapshot” of a ~ 25-hectare area frequented by Neanderthals during the Last Interglacial, with a focus on their exploitation of the pond terrapin Emys orbicularis. These data constitute the first evidence of turtle exploitation by Neanderthals north of the European mountain chains, beyond the Mediterranean basin. This Neumark-Nord record demonstrates that Last Interglacial foragers exploited a wide range of archaeologically visible resources available in this lake area, from small (~ 1 kg) pond terrapins up to and including the largest terrestrial mammals of the Pleistocene, straight-tusked elephants, with adult males weighing more than 10 tonnes. The abundance of intensively exploited medium- and large-sized mammals found alongside these Emys remains suggests that other variables than macronutrients per se played a role in the repeated harvesting of pond terrapins from these water bodies.
Neanderthal and anatomically modern human (AMH) occupations have long been hypothesized to differ in spatial organization, yet multisite quantitative comparisons remain scarce. Here we use cluster modeling to quantify four spatial metrics (parent intensity κ, cluster scale s, cluster strength ϕ, and sibling probability p) from 21 Middle Paleolithic (MP) and Upper Paleolithic (UP) assemblages, spanning cave and open-air contexts. We combine these metrics into two principal axes-compactness vs. dispersion and clustering scale-and assess their relationship to cultural period, site context, hearth counts, and functionality through regression analyses. Our results show that cultural affiliation is the strongest and most consistent predictor: UP sites are significantly more compact than MP ones, independent of context or occupation features. This pattern suggests that AMH groups structured their living spaces in more spatially constrained and segregated ways, with implications for the evolution of camp organization, activity planning, and social cohesion. At the same time, we find marked variability within MP sites, with some Neanderthal occupations approaching the spatial compactness typical of UP sites. These results provide the first multivariate evidence that while broad differences exist between Neanderthals and AMHs, elements of 'modern' spatial behavior also emerged among Neanderthals, pointing to a more complex and overlapping processes of social and spatial organization in the Late Pleistocene.
The emerging picture of hominin evolution is one of complexity, population structure, and gene flow, which recent genomic inference approaches have begun to resolve. Among these are methods based on two-locus statistics, which summarize information contained in genealogical correlations between linked loci. Although the inclusion of ancient samples could provide increased power to distinguish between competing models, these methods typically rely on large samples from present-day populations, and it remains challenging to apply them to ancient DNA (aDNA), which is sparsely sampled, unphased, and time-stratified. Here we develop an inference framework based on a set of multi-population two-locus statistics that are applicable to aDNA because they can be estimated from single unphased diploid genomes. We connect these statistics to an existing system of two-locus summaries and use them to model divergence and gene flow among populations represented by seven ancient hominin individuals and one contemporary human. We infer a demographic model with two episodes of gene flow from early anatomically modern humans (AMH) to Neanderthals and an introgression from an unsampled hominin lineage to Denisovan ancestors, broadly consistent with previous work. We also learn parameters of ancient Eurasian AMH population structure, reinforcing previous findings that early European farmers traced a large fraction of their ancestry to a lineage which split early from other non-African AMH and received little or no introgression from Neanderthals. Using both simulation and empirical data, we show that accurately estimating parameters associated with multiple gene flow episodes requires their joint inference due to their correlated effects on diversity.
N-acetyltransferase 1 (NAT1) is an enzyme that acetylates certain drugs and carcinogens and is involved in folate metabolism. Some NAT1 variants are associated with susceptibility to cancer and birth defects. We show that while nearly all present-day humans carry a valine residue at position 149 and a serine residue at position 214, some carry the ancestral amino acids isoleucine and alanine at these positions because of gene flow from Neanderthals. Neither substitution affects enzymatic activity. However, replacing the serine residue, a known phosphorylation site, with a phosphomimetic aspartate reduces NAT1 activity by ~60%. Phosphorylation at this site may allow NAT1 to be down-regulated during pregnancy to reduce birth defect risk, while maintaining high activity in adults for xenobiotic clearance. This regulatory site is a molecular feature that differs between modern humans and Neanderthals.
Neanderthals of Central-Eastern Europe are well documented by a wealth of archaeological sites, but thus far they remain poorly represented by both fossil and genetic data.1,2,3 At Stajnia Cave (Poland), nine Neanderthal teeth have now been integrated into a single high-resolution study combining morphological assessment, radiocarbon dating, and complete mitochondrial (mt) genome sequencing. We report eight new mitogenomes, including from four never-before-analyzed teeth, that resolve a minimum of seven, and possibly eight individuals. Three of the specimens share identical mitochondrial DNA (mtDNA), indicating that they are either from the same or maternally related individuals. Molecular branch shortening estimates place all samples in marine isotopic stage (MIS) 5, with point estimates of ∼119,700-92,498 years ago, making this the oldest multi-individual Neanderthal genetic assemblage yet characterized in Central Europe. Other Neanderthals with similar haplotypes were present in southeastern France, Iberia, and the Caucasus, suggesting this mt lineage might have been widespread across Europe before being replaced with the mtDNA of the "late Neanderthal type." Our analysis of the Stajnia fossils positions Central-Eastern Europe not as a peripheral fringe, but as a pivotal area for tracing Neanderthal geographic distribution.
Background/Objectives: Along the Tyrrhenian coast of central Italy, multilayered caves have yielded significant Neanderthal-era human remains. Recent excavations at Guattari Cave uncovered hominin fossils dated to approximately 66-65 ka, revealing a population with notable morpho-anatomical variability exhibiting both plesiomorphic (primitive) and autapomorphic (derived) traits. Methods: Here we present detailed morphometric and comparative analyses of cranial, dental, and postcranial remains, demonstrating affinities with Homo erectus (sensu stricto [s.s.] and lato [s.l.]), Proto-Neanderthals, classical Neanderthals, and Homo sapiens. Results: These findings indicate notable morpho-anatomical variability among the Guattari Cave hominin remains, with affinities to multiple hominin lineages during the Middle and Late Pleistocene. Pleistocene. Conclusions: The Guattari Cave assemblage thus contributes to our understanding of Eurasian hominin diversity and evolutionary dynamics, highlighting the Mediterranean as a region of interest for studying the phyletic continuity and diversity preceding modern humans.