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.
Preeclampsia and eclampsia, unique to human reproduction, represent the first disease documented in written history over 5000 years ago, yet their etiology remains elusive in 2026. These disorders, exclusive to Homo sapiens among 4300 mammal species, may have posed an even greater reproductive challenge to Neanderthals, potentially contributing to their poor fecundity. Arising from incomplete deep trophoblast invasion into maternal spiral arteries, essential for nourishing the energy-demanding fetal brain, they lead to placental insufficiency and fetal growth restriction (FGR). In humans, eclampsia (grand mal seizures) occurs naturally in ∼1 % of pregnancies, while preeclampsia affects 2-8 %, with untreated cases carrying high maternal and fetal mortality. Predominantly affecting primiparas and multiparas with a new partner ("primipaternity"), early-onset preeclampsia (EOP; delivery <34 weeks) results from failed maternal immune tolerance to paternal antigens, causing partial fetal rejection and inadequate artery remodeling. This manifests as FGR with or without maternal syndrome. Critically, humans evolved a protective mechanism decoupling maternal preeclampsia from ∼75 % of placental FGR cases, averting life-threatening complications. Without this safeguard, preeclampsia rates could soar to 10-20 %, with eclampsia at 4-5 %, severely impeding reproductive success. Neanderthals, sharing deep hemochorial placentation but possibly lacking this adaptation due to genetic divergences (e.g., imprinted genes, KIR-HLA interactions, PIEZO1 variants), likely suffered higher incidences, exacerbating demographic vulnerabilities like small populations and inbreeding. This hypothesis bridges a gap in paleoanthropology: preeclampsia, the principal human reproductive complication, is never cited by anthropologists as possible explanation of the well-known low fecundity rates in Neanderthals communities.
Sex biases in admixture and other demographic processes are recurrent features throughout human evolution. For admixture between Neanderthals and anatomically modern humans (AMHs), sex bias has been proposed as an explanation for the relative lack of Neanderthal ancestry in modern human X chromosomes compared with that in modern human autosomes. By observing a 62% relative excess of AMH ancestry in Neanderthal X chromosomes, we characterized the interbreeding between the two groups as predominantly male Neanderthals with female AMHs. Analytic and numerical modeling presents mate preference as a more parsimonious cause of the sex bias than purely demographic processes with differential patterns of male and female migration.
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.
There remains debate about the pyrotechnical capabilities of Neanderthals. Evidence of fire has been found at many Middle Palaeolithic sites, widely accepted to be associated with Neanderthals. However, multiple Neanderthal sites show a marked decrease in evidence for fire use during colder periods. This counterintuitive pattern was explained by the possibility that some Neanderthal groups were unable to create fire at will and relied on wildfire. Here, we evaluate the plausibility of this "wildfire hypothesis" through formal modeling. We computed the probability of a group of Neanderthals losing campfire-making skills due to cultural loss. The EMBERS model codes four empirically motivated mechanisms of skill loss: variability in use, period in between uses, memory decay and number of experts. Our results indicate that losing the ability to use wildfire was more likely than retaining it for most of our parameter values within reasonable ranges. Significantly, demography, in the form of expert numbers, was the least significant mechanism of loss. The rate of memory loss at group level, and intervals between uses were markedly more important than demography. Variability in time between uses was by far the strongest driver of loss. These results, linked with the estimated climatic, mnemonic, and demographic conditions for the Neanderthals' occupation of Europe in cold periods, support the plausibility of the wildfire hypothesis. Our results also highlight the need to pay more attention to cultural loss as a factor in cultural evolution. Our modeling demonstrates the feasibility of the controversial hypothesis that some European Neanderthal groups were unable to create fire at will and instead relied on wildfire to start their campfires.
Located in the northernmost part of Central Asia, the western foothills of the Altai Mountains (Western Siberia) represent to date the easternmost known boundary of Neanderthal distribution, far from their main cultural areas currently known in Western Eurasia. This geographic situation suggests the possibility of distinct cultural and biological traits in Altai Neanderthals. In this region, Chagyrskaya Cave contains the most substantial paleoanthropological collection, with 75 remains, including 20 craniodental elements attributed to at least eight individuals of varying ages (22 permanent teeth and four deciduous teeth), dating to between approximately 59 and 51 ka BP. Previous paleogenetic analyses suggest several individuals from this site are closely related. Our study is the first to comprehensively analyze the morphology of the entire set of dentognathic elements. In this study, we document the phenotypic variability of the Chagyrskaya's individuals by examining the dimensions and proportions of the crown and root tissues, the nonmetric traits of the outer enamel surface, and the enamel-dentine junction of the 26 teeth from this site and by comparing them to published data of both fossil and more recent material. Furthermore, we explore aspects related to their lifestyle and behavior describing the antemortem lesions affecting their dentognathic elements. Our results show that the dental traits of these human remains fall within the known Neanderthal phenotypic variability while also presenting certain specificities, the origins of which we discuss. In addition, the identification of several lesions on some of these fossils allows us to document their oral health and the use of their teeth for paramasticatory activities.
A pair of studies illuminates these humans' long, hardscrabble existence.
Human chromosome 3p21.31 variants introgressed from Neanderthals have been associated with a higher risk of developing a severe form of COVID-19. These Neanderthal DNA variants would regulate the expression of several genes, including LZTFL1 (implicated in the epithelial-mesenchymal transition) and proinflammatory chemokine receptors. We studied three introgressed haplotypes in patients who developed critical COVID-19 (N = 446; 82 deaths), less severe non-critical COVID-19 (N = 552), and population controls (N = 500) from the region of Asturias, Northern Spain. All the participants were genotyped for six single nucleotide polymorphisms that defined the three 3p21.31 haplotypes. For the haplotype in the LZTFL1 gene, the total patients were significantly higher frequency carriers of the Neanderthal variant compared to controls (24% vs. 17%; p < 0.05, OR = 1.53, 95% CI = 1.16-2.01). Multiple logistic regression showed that critical COVID-19 was independently associated with male sex, hypertension, dyslipaemia, and the introgressed LZTFL1 haplotype (p = 0.006). The frequency of these introgressed genotypes did not differ between normotensives and normolipaemics in the two patient groups but was significantly increased among hypertensives (p = 0.003) and dyslipaemics (p = 0.001). In our population, the 3p21.31 haplotypes introgressed from Neanderthals were associated with increased risk of critical COVID-19, and the risk effect was higher among patients with hypertension and dyslipaemia.
暂无摘要（点击查看详情）
暂无摘要（点击查看详情）
暂无摘要（点击查看详情）
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.
In this study we construct lists of candidate genes for articulate language. Analysis of coding regions of over 100 candidate genes for the effects of natural selection (directional episodic selection and relaxed/intensified selection) in the various lineages of primates (thirty-four nonhuman primate species, plus Homo sapiens Neanderthals and Denisovans) revealed a burst of altered selection effects on neural genes at the node leading to the Homo sapiens-Neanderthal-Denisova triad, followed by bursts of selection effects on neural genes related to language in both the Denisovan and Neanderthal lineages. Those latter increases in involvement of neural genes in Neanderthals and Denisovans can be contrasted with the missing or slight response to selection on those same genes in the H. sapiens lineage. The genes involved in these bursts can mostly be classified as involved in synapse structure and maintenance. We develop a hypothesis for how synaptic efficiency could be related to language acquisition in these lineages.
What genes and regulatory sequences critically differentiate modern humans from apes and archaic humans, which share highly similar genomes but show distinct phenotypes, has puzzled researchers for decades. Previous studies examined species-specific protein-coding genes and related regulatory sequences, revealing that birth, loss, and changes in these genes and sequences drive speciation and evolution. However, investigations of species-specific lncRNA genes and related regulatory sequences, which regulate substantial genes, remain limited. We identified human-specific (HS) lncRNAs from GENCODE-annotated human lncRNAs, predicted their DNA-binding domains (DBDs) and DNA-binding sites (DBSs), analyzed DBS sequences in modern humans (CEU, CHB, and YRI), archaic humans (Altai Neanderthals, Denisovans, and Vindija Neanderthals), and chimpanzees, and investigated how HS lncRNAs and their DBSs have influenced gene expression in archaic and modern humans. Our results suggest that these lncRNAs and DBSs have substantially reshaped gene expression, and this reshaping has evolved continuously from archaic to modern humans, enabling humans to adapt to new environments and lifestyles, promoting brain evolution, and resulting in cross-population differences. The parallel analysis of gene expression in GTEx tissues by HS transcription factors (TFs) and their DBSs indicates that HS lncRNAs have reshaped gene expression in the brain more significantly than HS TFs.
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.
Neanderthal endocrania are different in shape, though slightly larger in size than modern humans on average. These shape differences have long been used to suggest Neanderthals differed cognitively from modern humans, e.g., by having inferior linguistic/symbolic ability, poorer executive function, and/or smaller episodic and working memory capacity. However, whether the morphological differences in their brains inferred from their endocrania indicate truly meaningful cognitive/behavioral differences-with real evolutionary implications-is not clear. Recent work using deformation mapping techniques suggested there had been significant brain differences between Neanderthal and the anatomically modern Homo sapiens (amHs) that were contemporary with them. However the inferred differences were not put into the context of modern human populational variation in brain anatomy, which is known to be substantial. We estimate MRI brain region volumes in two modern population samples (US and Chinese) using deformation mapping and find similar sized differences to those reported for Neanderthal vs. amHs, with 9 of 13 brain regions showing larger absolute differences for US vs. Chinese samples as compared to reported Neanderthal vs. amHs differences. To the extent that brain anatomy differences indicate cognitive differences, this suggests that cognitive differences between Neanderthals and amHs would have comfortably fit within the range found among modern human populations-which are generally not considered evolutionarily significant. In fact, the endocranial differences between Neanderthal and their contemporaries predict cognitive difference effect sizes of only 0.14 SDs or less. This undermines the suggestion that Neanderthal replacement occurred because of cognitive limitations.
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.
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.
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.