Neanderthal Populations Research Articles

An ecological niche shift for Neanderthal populations in Western Europe 70,000\xa0years ago

Middle Paleolithic Neanderthal populations occupied Eurasia for at least 250,000 years prior to the arrival of anatomically modern humans. While a considerable body of archaeological research has focused on Neanderthal material culture and subsistence strategies, little attention has been paid to the relationship between regionally specific cultural trajectories and their associated existing fundamental ecological niches, nor to how the latter varied across periods of climatic variability. We examine the Middle Paleolithic archaeological record of a naturally constrained region of Western Europe between 82,000 and 60,000 years ago using ecological niche modeling methods. Evaluations of ecological niche estimations, in both geographic and environmental dimensions, indicate that 70,000 years ago the range of suitable habitats exploited by these Neanderthal populations contracted and shifted. These ecological niche dynamics are the result of groups continuing to occupy habitual territories that were characterized by new environmental conditions during Marine Isotope Stage 4. The development of original cultural adaptations permitted this territorial stability.

Nubian Levallois technology associated with southernmost Neanderthals

Neanderthals occurred widely across north Eurasian landscapes, but between ~ 70 and 50 thousand years ago (ka) they expanded southwards into the Levant, which had previously been inhabited by Homo sapiens. Palaeoanthropological research in the first half of the twentieth century demonstrated alternate occupations of the Levant by Neanderthal and Homo sapiens populations, yet key early findings have largely been overlooked in later studies. Here, we present the results of new examinations of both the fossil and archaeological collections from Shukbah Cave, located in the Palestinian West Bank, presenting new quantitative analyses of a hominin lower first molar and associated stone tool assemblage. The hominin tooth shows clear Neanderthal affinities, making it the southernmost known fossil specimen of this population/species. The associated Middle Palaeolithic stone tool assemblage is dominated by Levallois reduction methods, including the presence of Nubian Levallois points and cores. This is the first direct association between Neanderthals and Nubian Levallois technology, demonstrating that this stone tool technology should not be considered an exclusive marker of Homo sapiens.

Erratum to “Palaeoecological context for the extinction of the Neanderthals: A small mammal study of Stratigraphic Unit V of the El Salt site, Alcoi, eastern Spain” [Palaeogeography, Palaeoclimatology, Palaeoecology 530 (2019) 163–175

Abstract El Salt is an important reference site for understanding the extinction of Neanderthal populations in the eastern Iberian Peninsula during MIS 3. In this paper, we describe the small mammal assemblage from Stratigraphic Unit V, the youngest unit with evidence of human presence, based on nearly 1300 specimens. A total of seven rodents (Microtus arvalis, Microtus duodecimcostatus, Microtus cabrerae, Sciurus vulgaris, Arvicola sapidus, Eliomys quercinus and Apodemus sylvaticus), three insectivores (Talpa occidentalis, Crocidura sp., Sorex sp.) and one lagomorph (Oryctolagus cf. cuniculus) were identified. Palaeocological analyses point to drier conditions in this part of the stratigraphic sequence, supporting the hypothesis that an aridification scenario may have played a role in the extinction of the Neanderthal groups inhabiting this region of the Iberian Peninsula.

Каменные индустрии ранних этапов среднего палеолита стоянки Дарвагчай-Залив-4 (исследования 2021 года)

The territory of Primorsky Dagestan is one of the places on the Eurasian continent, where a constant and massive presence of Neanderthal populations in antiquity was noted. In particular, this is evidenced by a large number of Middle Paleolithic sites in a relatively small territory of the Darvagchay geoarchaeological region (Southeastern Dagestan), discovered and studied in the last decade. This article is devoted to the results of the latest studies of the Middle Paleolithic complexes of the Darvagchay-Zaliv-4 site, the materials of which belong to the early stages of the Middle Paleolithic and significantly supplement the evidence material of this cultural-chronological period in the territory of the North-Eastern Caucasus. In the field season of 2021, two sets of artifacts related to different stages of the Middle Paleolithic were discovered at the site. The age of the geological horizons, where they were discovered, was previously based on the results of absolute dating by OSL. The stone industry of the first complex is characterized by the Levallois technique and a typical Middle Paleolithic toolkit; artifacts were located in horizon 1c. Chronologically, the time of its existence is determined by the period of the Riess-Wurm (Eem, Mikulino) interglacial, in the range of 120-110 ka BP (MIS 5e). Artifacts of another complex, discovered at the site for the first time, were found in the middle part of Layer 2 (light loam, loess). Aeolian-deluvial processes played a leading role in sedimentation here. The technical and typological characteristics of a small collection of stone products make it possible to attribute it to the earliest stages of the Middle Paleolithic. The age of the culture-bearing horizon was based on a series of the OSL dates in the range of 200-180 ka BP (MIS 8).

Depositional processes and environmental settings in rock shelters: the case of the prehistoric Oscurusciuto site (Southern Italy)

AbstractClastic successions in rock shelters commonly host important archaeological findings, especially of prehistoric and protostoric times. The understanding of depositional and post-depositional processes in these environments is crucial to understand the lifestyle settings of humans, as well as the reliability of archaeological data obtained during excavations. Rock shelters are genetically related to caves, but while depositional processes in caves are generally well known, less information is available concerning the depositional processes active in rock shelters.This paper tries to contribute to this issue, describing the sedimentary succession exposed at the Oscurusciuto rock shelter (Ginosa, Southern Italy). This is one of the most important Middle Palaeolithic sites of the Italian peninsula, and its sedimentary infill hosts witnesses of very late Neanderthal populations that lived in Italy just before their total decline and complete replacement by Modern Humans (MH).This work presents the results of a sedimentologically based study integrated with an ichnological study of selected beds. The combination of these two methodologies allowed us to: (i) define the main depositional processes active in the rock shelter environment, as well as the relationships between different processes; (ii) discuss the meaning of peculiar ichnofabrics recognized at the site, and (iii) discuss the meaning of structureless strata at an archeological site in the framework of human trampling vs bioturbation as a cause of the obliteration of primary sedimentary structures.

The early Aurignacian dispersal of modern humans into westernmost Eurasia

Documenting the first appearance of modern humans in a given region is key to understanding the dispersal process and the replacement or assimilation of indigenous human populations such as the Neanderthals. The Iberian Peninsula was the last refuge of Neanderthal populations as modern humans advanced across Eurasia. Here we present evidence of an early Aurignacian occupation at Lapa do Picareiro in central Portugal. Diagnostic artifacts were found in a sealed stratigraphic layer dated 41.1 to 38.1 ka cal BP, documenting a modern human presence on the western margin of Iberia ∼5,000 years earlier than previously known. The data indicate a rapid modern human dispersal across southern Europe, reaching the westernmost edge where Neanderthals were thought to persist. The results support the notion of a mosaic process of modern human dispersal and replacement of indigenous Neanderthal populations.

New perspectives on Neanderthal dispersal and turnover from Stajnia Cave (Poland)

The Micoquian is the broadest and longest enduring cultural facies of the Late Middle Palaeolithic that spread across the periglacial and boreal environments of Europe between Eastern France, Poland, and Northern Caucasus. Here, we present new data from the archaeological record of Stajnia Cave (Poland) and the paleogenetic analysis of a Neanderthal molar S5000, found in a Micoquian context. Our results demonstrate that the mtDNA genome of Stajnia S5000 dates to MIS 5a making the tooth the oldest Neanderthal specimen from Central-Eastern Europe. Furthermore, S5000 mtDNA has the fewest number of differences to mtDNA of Mezmaiskaya 1 Neanderthal from Northern Caucasus, and is more distant from almost contemporaneous Neanderthals of Scladina and Hohlenstein-Stadel. This observation and the technological affinity between Poland and the Northern Caucasus could be the result of increased mobility of Neanderthals that changed their subsistence strategy for coping with the new low biomass environments and the increased foraging radius of gregarious animals. The Prut and Dniester rivers were probably used as the main corridors of dispersal. The persistence of the Micoquian techno-complex in South-Eastern Europe infers that this axis of mobility was also used at the beginning of MIS 3 when a Neanderthal population turnover occurred in the Northern Caucasus.

Homo neanderthalensis and the evolutionary origins of ritual in Homo sapiens.

There is a large, if disparate, body of archaeological literature discussing specific instantiations of symbolic material culture and the possibility of ritual practices in Neanderthal populations. Despite this attention, however, no single synthesis exists that draws upon cognitive, psychological and cultural evolutionary theories of ritual. Here, we review the evidence for ritual-practice among now-extinct Homo neanderthalensis, as well as the necessary cognitive pre-conditions for such behaviour, in order to explore the evolution of ritual in Homo sapiens. We suggest that the currently available archaeological evidence indicates that Neanderthals may have used 'ritualization' to increase the successful transmission of technical knowledge across generations-providing an explanation for the technological stability of the Middle Palaeolithic and attesting to a survival strategy differing from near-contemporary H. sapiens. This article is part of the theme issue 'Ritual renaissance: new insights into the most human of behaviours'.

Inner morphological and metric characterization of the molar remains from the Montmaurin-La Niche mandible: The Neanderthal signal

Here, we present a metric and morphological study of the molar remains from the Montmaurin-La Niche mandible by means of microcomputed tomography. According to the last analysis, based on the combination of geomorphological and paleontological data, the level bearing this human mandible probably corresponds to the marine isotope stages (MIS) 7. These data place the Montmaurin-La Niche in a chronologically intermediate position between the Neanderthals and the Middle Pleistocene fossils (e.g., Sima de los Huesos, la Caune de l’Arago). A recent study has revealed that while the mandible is more closely related to the Early and Middle Pleistocene African and Eurasian populations, the morphology of the outer enamel surfaces of its molars is typical of the Neanderthal linage. The data presented here are in line with this finding because the morphology of the enamel-dentine junction of the molars is similar to that of Neanderthals, whereas the absolute and relative enamel thickness values (2D and 3D) are closer to those exhibited by some Early Pleistocene hominins. Moreover, the pulp cavity morphology and proportions are in concordance with the Neanderthal populations. Our results strengthen the hypothesis that the settlement of Europe could be the result of several migrations, at different times, originated from a common source population. Thus, the variability in the European Middle Pleistocene populations (e.g., Montmaurin, Sima de los Huesos, Arago, Mala Balanica) could indicate different migrations at different times and/or population fragmentation, without excluding the possible hybridization between residents and new settlers.

Analysis of Haplotypic Variation and Deletion Polymorphisms Point to Multiple Archaic Introgression Events, Including from Altai Neanderthal Lineage.

The time, extent, and genomic effect of the introgressions from archaic humans into ancestors of extant human populations remain some of the most exciting venues of population genetics research in the past decade. Several studies have shown population-specific signatures of introgression events from Neanderthals, Denisovans, and potentially other unknown hominin populations in different human groups. Moreover, it was shown that these introgression events may have contributed to phenotypic variation in extant humans, with biomedical and evolutionary consequences. In this study, we present a comprehensive analysis of the unusually divergent haplotypes in the Eurasian genomes and show that they can be traced back to multiple introgression events. In parallel, we document hundreds of deletion polymorphisms shared with Neanderthals. A locus-specific analysis of one such shared deletion suggests the existence of a direct introgression event from the Altai Neanderthal lineage into the ancestors of extant East Asian populations. Overall, our study is in agreement with the emergent notion that various Neanderthal populations contributed to extant human genetic variation in a population-specific manner.

Quantifying the potential causes of Neanderthal extinction: Abrupt climate change versus competition and interbreeding

Anatomically Modern Humans are the sole survivor of a group of hominins that inhabited our planet during the last ice age and that included, among others, Homo neanderthalensis, Homo denisova, and Homo erectus. Whether previous hominin extinctions were triggered by external factors, such as abrupt climate change, volcanic eruptions or whether competition and interbreeding played major roles in their demise still remains unresolved. Here I present a spatially resolved numerical hominin dispersal model (HDM) with empirically constrained key parameters that simulates the migration and interaction of Anatomically Modern Humans and Neanderthals in the rapidly varying climatic environment of the last ice age. The model simulations document that rapid temperature and vegetation changes associated with Dansgaard-Oeschger events were not major drivers of global Neanderthal extinction between 50 and 35 thousand years ago, but played important roles regionally, in particular over northern Europe. According to a series of parameter sensitivity experiments conducted with the HDM, a realistic extinction of the Neanderthal population can only be simulated when Homo sapiens is chosen to be considerably more effective in exploiting scarce glacial food resources as compared to Neanderthals.

Initial Upper Palaeolithic Homo sapiens from Bacho Kiro Cave, Bulgaria.

The Middle to Upper Palaeolithic transition in Europe witnessed the replacement and partial absorption of local Neanderthal populations by Homo sapiens populations of African origin1. However, this process probably varied across regions and its details remain largely unknown. In particular, the duration of chronological overlap between the two groups is much debated, as are the implications of this overlap for the nature of the biological and cultural interactions between Neanderthals and H.sapiens. Here we report the discovery and direct dating of human remains found in association with Initial Upper Palaeolithic artefacts2, from excavations at Bacho Kiro Cave (Bulgaria). Morphological analysis of a tooth and mitochondrial DNA from several hominin bone fragments, identified through proteomic screening, assign these finds to H.sapiens and link the expansion of Initial Upper Palaeolithic technologies with the spread of H.sapiens into the mid-latitudes of Eurasia before 45thousand years ago3. The excavations yielded a wealth of bone artefacts, including pendants manufactured from cave bear teeth that are reminiscent of those later produced by the last Neanderthals of western Europe4-6. These finds are consistent with models based on the arrival of multiple waves of H.sapiens into Europe coming into contact with declining Neanderthal populations7,8.

Cannibals in the forest: A comment on Defleur and Desclaux (2019)

Though rare in the archaeological record, cannibalism has been documented among many different hominin populations; however, the mechanisms driving such behaviors remains poorly understood. Recent research that evaluates the hypothesized Neanderthal nutritional cannibalism at the Middle Palaeolithic cave site of Baume Moula-Guercy (BMG) in Mediterranean France, is one case that, in our opinion, is not fully substantiated by the evidence put forth by Defleur and Desclaux (2019). Using site occurrence data from the Last Interglacial (MIS 5e), the biogeography of extinct reptile species, and cut mark taphonomic evidence, Defleur and Desclaux (2019) suggest that the Neanderthal remains found in Layer XV of the cave are the result of nutritional cannibalism. The authors argue that a regional collapse of Neanderthal populations was the result of a depletion of prey biomass, caused by a profound upheaval of local environmental conditions, thus causing the Neanderthals at BMG to resort to cannibalism. In this response paper, we attempt to show that the methods and materials used by Defleur and Desclaux 1) do not reflect the known number of Eemian-aged sites, 2) do not present sufficient evidence related to the environmental stress conditions at BMG that would warrant cannibalism, and 3) taphonomic evidence on Neanderthal remains does not strictly support the argument for nutritional cannibalism. Additional study of the BMG remains is necessary before ruling out other processes related to the post-mortem treatment of the Neanderthal corpses.

The study of the lower limb entheses in the Neanderthal sample from El Sidrón (Asturias, Spain): How much musculoskeletal variability did Neanderthals accumulate?

Entheses have rarely been systematically studied in the field of human evolution. However, the investigation of their morphological variability (e.g., robusticity) could provide new insight into their evolutionary significance in the European Neanderthal populations. The aim of this work is to study the entheses and joint features of the lower limbs of El Sidrón Neanderthals (Spain; 49 ka), using standardized scoring methods developed on modern samples. Paleobiology, growth, and development of both juveniles and adults from El Sidrón are studied and compared with those of Krapina Neanderthals (Croatia, 130 ka) and extant humans. The morphological patterns of the gluteus maximus and vastus intermedius entheses in El Sidrón, Krapina, and modern humans differ from one another. Both Neanderthal groups show a definite enthesis design for the gluteus maximus, with little intrapopulation variability with respect to modern humans, who are characterized by a wider range of morphological variability. The gluteus maximus enthesis in the El Sidrón sample shows the osseous features of fibrous entheses, as in modern humans, whereas the Krapina sample shows the aspects of fibrocartilaginous ones. The morphology and anatomical pattern of this enthesis has already been established during growth in all three human groups. One of two and three of five adult femurs from El Sidrón and from Krapina, respectively, show the imprint of the vastus intermedius, which is absent among juveniles from those Neanderthal samples and in modern samples. The scant intrapopulation and the high interpopulation variability in the two Neanderthal samples is likely due to a long-term history of small, isolated populations with high levels of inbreeding, who also lived in different ecological conditions. The comparison of different anatomical entheseal patterns (fibrous vs. fibrocartilaginous) in the Neanderthals and modern humans provides additional elements in the discussion of their functional and genetic origin.

Neanderthal-Denisovan ancestors interbred with a distantly related hominin.

Previous research has shown that modern Eurasians interbred with their Neanderthal and Denisovan predecessors. We show here that hundreds of thousands of years earlier, the ancestors of Neanderthals and Denisovans interbred with their own Eurasian predecessors-members of a "superarchaic" population that separated from other humans about 2 million years ago. The superarchaic population was large, with an effective size between 20 and 50 thousand individuals. We confirm previous findings that (i) Denisovans also interbred with superarchaics, (ii) Neanderthals and Denisovans separated early in the middle Pleistocene, (iii) their ancestors endured a bottleneck of population size, and (iv) the Neanderthal population was large at first but then declined in size. We provide qualified support for the view that (v) Neanderthals interbred with the ancestors of modern humans.

Archaeological evidence for two separate dispersals of Neanderthals into southern Siberia

Neanderthals were once widespread across Europe and western Asia. They also penetrated into the Altai Mountains of southern Siberia, but the geographical origin of these populations and the timing of their dispersal have remained elusive. Here we describe an archaeological assemblage from Chagyrskaya Cave, situated in the Altai foothills, where around 90,000 Middle Paleolithic artifacts and 74 Neanderthal remains have been recovered from deposits dating to between 59 and 49 thousand years ago (age range at 95.4% probability). Environmental reconstructions suggest that the Chagyrskaya hominins were adapted to the dry steppe and hunted bison. Their distinctive toolkit closely resembles Micoquian assemblages from central and eastern Europe, including the northern Caucasus, more than 3,000 kilometers to the west of Chagyrskaya Cave. At other Altai sites, evidence of earlier Neanderthal populations lacking associated Micoquian-like artifacts implies two or more Neanderthal incursions into this region. We identify eastern Europe as the most probable ancestral source region for the Chagyrskaya toolmakers, supported by DNA results linking the Neanderthal remains with populations in northern Croatia and the northern Caucasus, and providing a rare example of a long-distance, intercontinental population movement associated with a distinctive Paleolithic toolkit.

Volcanism and Settlement of the Northern Slope of the Central Caucasus in the Middle Paleolithic: New Data from Saradj-Chuko Grotto

The problem of the settlement of the territory of the Northern Caucasus in the Paleolithic and its dependence on volcanic activity factors, climate, and shift in ecological niches is very relevant in modern domestic and foreign studies. Previous research at Mezmaiskaya cave in the northwestern Caucasus revealed that volcanism significantly affected the Middle Paleolithic population of this region and the disappearance of the Neanderthal population. In 2016, the first stratified Middle Paleolithic site was discovered in the Elbrus region (Republic of Kabardino–Balkaria), on the northern slopes of the central Caucasus. This site was called Saradj-Chuko grotto. This region is fundamentally important for understanding the dynamics of the Northern Caucasus settlement in the Paleolithic because it is located at the crossroads between the northern and southern slopes. The only obsidian outcrop in the Northern Caucasus is located also here. Obsidian is a volcanic raw material that was highly valued by ancient people in that period and transported by Neanderthals hundreds of kilometers. We have investigated the chemical composition and morphology of minerals in 76 samples of the ZZ0 longitudinal section in Saradj-Chuko grotto. As a result, traces of volcanic activity were revealed in 2 (6A and 4) out of 11 layers. It has been found that volcanism in the Middle Paleolithic affected the settlement of the Elbrus region and most likely the entire northern slope of the central Caucasus. Specifically, layer 6A (where ash was found) overlaps layer 6B (the level of active habitation in the grotto). After the eruption recorded in layer 6A, the Neanderthals only occasionally visited the cave. Preliminary conclusions are made about a possible source of ash in these layers of Saradj-Chuko grotto. The materials were analyzed in a wide context of the currently known data on the presence of ashes in other sites of the Caucasus.

Fire and brief human occupations in Iberia during MIS 4: Evidence from Abric del Pastor (Alcoy, Spain)

There is a relatively low amount of Middle Paleolithic sites in Europe dating to MIS 4. Of the few that exist, several of them lack evidence for anthropogenic fire, raising the question of how this period of global cooling may have affected the Neanderthal population. The Iberian Peninsula is a key area to explore this issue, as it has been considered as a glacial refugium during critical periods of the Neanderthal timeline and might therefore yield archaeological contexts in which we can explore possible changes in the behaviour and settlement patterns of Neanderthal groups during MIS 4. Here we report recent data from Abric del Pastor, a small rock shelter in Alcoy (Alicante, Spain) with a stratified deposit containing Middle Palaeolithic remains. We present absolute dates that frame the sequence within MIS 4 and multi-proxy geoarchaeological evidence of in situ anthropogenic fire, including microscopic evidence of in situ combustion residues and thermally altered sediment. We also present archaeostratigraphic evidence of recurrent, functionally diverse, brief human occupation of the rock shelter. Our results suggest that Neanderthals occupied the Central Mediterranean coast of the Iberian Peninsula during MIS 4, that these Neanderthals were not undergoing climatic stress and they were habitual fire users.

Inbreeding, Allee effects and stochasticity might be sufficient to account for Neanderthal extinction.

The replacement of Neanderthals by Anatomically Modern Humans has typically been attributed to environmental pressure or a superiority of modern humans with respect to competition for resources. Here we present two independent models that suggest that no such heatedly debated factors might be needed to account for the demise of Neanderthals. Starting from the observation that Neanderthal populations already were small before the arrival of modern humans, the models implement three factors that conservation biology identifies as critical for a small population’s persistence, namely inbreeding, Allee effects and stochasticity. Our results indicate that the disappearance of Neanderthals might have resided in the smallness of their population(s) alone: even if they had been identical to modern humans in their cognitive, social and cultural traits, and even in the absence of inter-specific competition, Neanderthals faced a considerable risk of extinction. Furthermore, we suggest that if modern humans contributed to the demise of Neanderthals, that contribution might have had nothing to do with resource competition, but rather with how the incoming populations geographically restructured the resident populations, in a way that reinforced Allee effects, and the effects of inbreeding and stochasticity.

Provenance, modification and use of manganese-rich rocks at Le Moustier (Dordogne, France).

The use of colouring materials by Neanderthals has attracted a great deal of attention in recent years. Here we present a taphonomic, technological, chemical-mineralogical and functional analysis of fifty-four manganese rich lumps recovered during past and on-going excavations at the lower rockshelter of Le Moustier (Dordogne, France). We compare compositional data for archaeological specimens with the same information for twelve potential geological sources. Morphometric analysis shows that material from Peyrony’s excavations before the First World War provides a highly biased picture of the importance of these materials for Mousterian groups. These early excavations almost exclusively recovered large modified pieces, while Mn-rich lumps from the on-going excavations predominantly consist of small pieces, only half of which bear traces of modification. We estimate that at least 168 pieces were not recovered during early work at the site. Neanderthals developed a dedicated technology for processing Mn-rich fragments, which involved a variety of tools and motions. Processing techniques were adapted to the size and density of the raw material, and evidence exists for the successive or alternating use of different techniques. Morphological, textural and chemical differences between geological and archaeological samples suggest that Neanderthals did not collect Mn-rich lumps at the outcrops we sampled. The association and variability in Mn, Ni, As, Ba content, compared to that observed at the sampled outcrops, suggests that either the Le Moustier lumps come from a unique source with a broad variation in composition, associating Mn, Ni, As, Ba, or that they were collected at different sources, characterized either by Mn-Ni-As or Mn-Ba. In the latter case, changes in raw material composition across the stratigraphy support the idea that Neanderthal populations bearing different stone tool technologies collected Mn fragments from different outcrops. Our results favour a use of these materials for multiple utilitarian and symbolic purposes.